Genus Orestias (Cyprinodontiformes, Cyprinodontidae)

A TAXONOMIC REVISION OF THE ANDEAN KILLIFISH GENUS ORESTIAS (CYPRINODONTIFORMES, CYPRINODONTIDAE)

LYNNE R. PARENTI

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 178: ARTICLE 2 NEW YORK: 1984

A TAXONOMIC REVISION OF THE ANDEAN KILLIFISH GENUS ORESTIAS (CYPRINODONTIFORMES, CYPRINODONTIDAE)

LYNNE R. PARENTI Research Associate, Department ofIchthyology American Museum ofNatural History

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 178, article 2, pages 107-214, figures 1-72, tables 1-12 Issued May 9, 1984 Price: $8.40 a copy

Copyright ©) American Museum of Natural History 1984 ISSN 0003-0090 CONTENTS Abstract ...... 110 Introduction ...... 110 Acknowledgments ...... 113 Abbreviations ...... 114 Note on Materials and Methods ...... 115 Relationships of Killifishes of the Tribe Orestiini ...... 116 Phylogenetic Analysis ...... 122 Squamation and Neuromast Pattern ...... 123 Hyobranchial Apparatus ...... 126 Fins ...... 129 Chromosomes ...... 130 Sexual Dimorphism and Dichromatism ...... 131 Jaw and Jaw Suspensorium ...... 133 Skull ...... 134 Meristic Characters ...... 135 Morphometric Characters ...... 146 Explanation of Synapomorphy Diagrams ...... 150 Key to Orestias Species ...... 160 Systematic Accounts ...... 165 Genus Orestias Valenciennes ...... 165 Orestias cuvieri Valenciennes ...... 167 Orestias pentlandii Valenciennes ...... -1-68 Orestias ispi Lauzanne ...... 169 Orestiasforgeti Lauzanne ...... 170 Orestias mulleri Valenciennes ...... 171 Orestias gracilis, New Species ...... 172 Orestias crawfordi Tchernavin ...... 173 Orestias tutini Tchemavin ...... 174 Orestias incae Garman ...... 174 Orestias luteus Valenciennes ...... 175 Orestias rotundipinnis, New Species ...... 176 Orestiasfarfani, New Species ...... 178 Orestias albus Valenciennes ...... 179 Orestias olivaceus Garman ...... 180 Orestias silustani Allen ...... 181 Orestias agassii Valenciennes ...... 182 Orestias empyraeus Allen ...... 185 Orestiasfrontosus Cope ...... 186 Orestias polonorum Tchemavin ...... 188 Orestias elegans Garman ...... 188 Orestias jussiei Valenciennes ...... 189 Orestias puni Tchernavin ...... 189 Orestias parinacotensis Arratia ...... 190 Orestias laucaensis Arratia ...... 190 Orestias tschudii Castelnau ...... 191 Orestias gymnotus, New Species ...... 192

Orestias hardini, New Species ...... 193

Orestias ctenolepis, New Species ...... 194

Orestias ascotanensis, New Species...... 196

Orestias richersoni, New Species ...... 197

Orestias multiporis, New Species ...... 198

Orestias mundus, New Species ...... 199

Orestias ututo, New Species ...... 200

Orestias gilsoni Tchemavin...... 201

Orestias taquiri Tchernavin...... 201

Orestias mooni Tchemavin ...... 202 Orestias uruni Tchemavin...... 203

Orestias minimus Tchernavin...... 203

Orestias minutus Tchemavin ...... 204

Orestias tchernavini Lauzanne ...... 205

Orestias imarpe, New Species...... 205

Orestias tomcooni, New Species...... 206

Orestias robustus, New Species...... 207

?Orestias cuvieri X pentlandii ...... 208

Orestias rospigliosii Eigenmann and Allen...... 209

Orestias pentlandii var. fuscus Garman ...... 209

Summary ...... 209

Literature Cited ...... 211 ABSTRACT The ichthyofauna of the high Andes is assumed is a unique squamation and head pore pattern to be of low taxonomic diversity. Only three fish characterized by a prominent lyre-shaped arrange- genera, the killifish Orestias and the catfishes As- ment of minute neuromasts and a prominent me- troblepus and Trichomycterus include species en- dian dorsal ridge of scales running from the top demic to the Altiplano, the high-altitude plateau of the head to the dorsal fin origin. between the eastern and western slopes ofthe cen- Primary characters used to distinguish Orestias tral Andes. The atherinid Basilichthys is known species are overall body shape and squamation doubtfully from the Altiplano. pattern. Diversity of form is marked. There are In the last revision ofthe genus Orestias, Tcher- large, troutlike midwater predators, such as 0. cu- navin (1 944a) recognized 20 species and five sub- vieri, diminutive inshore species such as 0. mi- species. This small number was taken as evidence nutus and 0. minimus, and very wide-headed and of the low diversity of fish of the Titicaca Basin. wide-bodied species, with large, thick granulated However, in the past two years, five new Orestias scales, such as 0. luteus and 0. albus. species have been described, three from Lago Ti- The diversity among Orestias species in Lago ticaca (Lauzanne, 1981) and two from the Alti- Titicaca, the large, high-altitude lake of the Alti- plano of northern Chile (Arratia, 1982). With the plano of Peru and Bolivia, has led to the appli- benefit of large recent collections, 43 species of cation of the term "species flock" to this group of Orestias are recognized in the present revision, 14 lacustrine killifish. However, the Orestias of Ti- being described as new (0. gracilis, 0. rotundi- ticaca do not form a monophyletic group, and it pinnis, 0. farfani, 0. gymnotus, 0. hardini, 0. is recommended that the term "species flock" not ctenolepis, 0. ascotanensis, 0. richersoni, 0. mul- be used for the killifishes limited to the lake. tiporis, 0. mundus, 0. ututo, 0. imarpe, 0. tom- The known range of Orestias extends from cooni, and 0. robustus). northern Peru to northern Chile. It is expected that Seven synapomorphies described here define the number will rise with additional collecting, Orestias as a monophyletic group: (1) pelvic fins because collections are lacking from many of the and fin girdle are absent; (2) vomer is absent; (3) small, isolated lakes of the Altiplano; also, single middle anal and middle dorsal fin radials are car- specimens of apparently new species are known, tilaginous, rather than ossified; (4) bony anterior yet remain undescribed. and posterior ceratohyal are separated ventrally The hypothesis of a close relationship between by a large gap filled with cartilage; (5) first postclei- Orestias and the Anatolian cyprinodonts is re- thrum is absent; (6) anguloarticular lacks a ventral viewed. extension parallel to the retroarticular; and (7) there

INTRODUCTION The killifish genus Orestias (Cyprinodon- fish genera with some species endemic to the tiformes, Cyprinodontidae) is endemic to the Altiplano of the central Andes. high-altitude lakes and tributary streams of More than halfofthe 43 species of Orestias the Peruvian, Bolivian, and Chilean Andes. are endemic to the closed Titicaca Basin, and In the present revision the genus is consid- of those, 23 are known only from Lago Ti- ered to comprise 43 species. The range of ticaca, the large (8100 km2 surface area), high- Orestias (fig. 1) extends from Ancash Prov- altitude (3803 m) lake of the Altiplano, sit- ince, northern Peru to Antofagasta Province, uated between lat. 15°S and 17°S and between northern Chile. The distribution is coinci- long. 68°W and 7 1°W (with the midpoint of dent only in part with the catfish genera As- the lake at approximately 1 5045'S and troblepus and Trichomycterus (see Arratia, 69°25'W) in Peru and Bolivia (Richerson, 1982 and Arratia et al., 1978) and the sil- Widmer, and Kittel, 1977). verside genus Basilichthys,' the only other In the last revision of Orestias, Tchernavin (1 944a) formally recognized 20 species and I The genus Basilichthys is known as an Altiplano en- five subspecies. Recently, three more species demic from just one specimen, 17.5 mm SL, FMNH have been described from Lago Titicaca 94184, collected by W. R. Allen, 1918-1919, from a (Lauzanne, 1981), and two from the Alti- pool below Acora, Peru, a village near Chucuito, near plano of northern Chile (Arratia, 1982). In Bahia de Puno (fig. 22). the present paper, 14 species are described as 110 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS illl

FIG. 1. Limits of the known range of the genus Orestias, cross-hatched and outlined by a solid line. new. The majority of material used in the (1876, 1895) greatly increased our knowledge present revision was collected in five expe- of the Lago Titicaca ichthyofauna. ditions. The genus Orestias was first diag- The third major expedition which estab- nosed by Valenciennes (1839) in his report lished the abundance of species throughout on the collection by Pentland of a unique most ofthe wide-ranging distributional limits group of species from Lago Titicaca. of Orestias, including the Titicaca, Urubama, The second major expedition was by Alex- and Huallaga basins, was that by C. H. and ander Agassiz and Samuel W. Garman ofthe A. Eigenmann and W. R. Allen of Indiana Museum of Comparative Zoology, Harvard University. In 1918 and 1919 their expedi- University. Revisions of Orestias by Garmarn tion collected Orestias from Lago Junin (Lago 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 1 78

Chinchaycocha) in northern Peru, to Lago and continue to the present day in an effort Ascot'an in northern Chile. In part, this ma- to create a commercial trout fishery. The in- terial formed the basis of a revision of Or- troductions were successful and the fishery estias by Allen, in Eigenmann and Allen existed on the lake until 1969 (Laba, 1979). (1942). However, Allen did not use most of Reasons for the collapse of the commercial the more than 10,000 specimens of Orestias fishery, economical, sociological, and biolog- collected on this expedition in his revision. ical are varied (see Everett, 1971, 1973; Laba, They were distributed, many unsorted and 1979). However, the presence ofexotic species uncatalogued, to the collections of the Cali- has had a negative biological effect on the fornia Academy of Sciences and the Field native Orestias. The largest Orestias species, Museum ofNatural History. Representatives 0. cuvieri, has not been obtained in recent of several species were also deposited in the decades and is presumed extinct. The threats National Museum ofNatural History and the to Orestias come from the trout, and the sil- University ofMichigan, Museum ofZoology. verside Basilichthys bonairensis introduced The Percy Sladen Expedition of 1937 added in the 1950s (Laba, 1979). greatly to our knowledge ofthe Lago Titicaca Reports that the genus Orestias is nearly ichthyofauna (see Transactions of the Lin- extinct in Titicaca (e.g., Gery, 1969; Vill- nean Society, 1939-1955, for a comprehen- wock, 1972) are exaggerated, however. Large sive report on the Percy Sladen Expedition). collections from Titicaca made by Coon and This material at the British Museum (Natural a French team from the Office de la Re- History) formed nearly the entire basis of cherche Scientifique et Technique Outre Mer Tchernavin's (1944a) revision of Orestias, as (O.R.S.T.O.M.) reveal that many species are well as his revision of Trichomycterus still abundant, and that new species are pres- (Tchernavin, 1944b). ent (Lauzanne, 1981; present revision). Per- Large collections of Orestias, primarily haps more important, from a biogeograph- from Lago Titicaca, made in 1979 by Tom ical perspective, recent collecting has greatly Coon, then with the University of California expanded the known distributional limits of at Davis, are reviewed in the present study. the genus. Tim Hardin, an ecologist working The material has been deposited in the Amer- through Colorado State University, in 1979 ican Museum of Natural History, National discovered Orestias in three small lakes sit- Museum ofNatural History, Smithsonian In- uated northeast of Lima. This discovery not stitution, and the British Museum (Natural only revealed two new species, but also the History). possibility that Orestias species may be found Various geological hypotheses have been in the numerous small lakes that characterize proposed concerning the age ofLago Titicaca the Andes south of Ecuador. This revision of (Moon, 1939; Newell, 1949; James, 1971, Orestias should, therefore, be considered pre- 1973). The isolation of the Altiplano from liminary, in that it attempts to summarize ocean drainage systems is estimated to have the known diversity of Orestias, with the occurred in the Miocene, and the formation knowledge that searches for a limit to its range, of Titicaca by Plio-Pleistocene times (James, and assessment ofabundance throughout the 1973). Today, Titicaca is composed of three range, have still to be made. Emphasis is major sections (fig. 22): Lago Grande, a large placed on morphology; forexample, a descrip- steep-sided, deep (to 281 m) basin, and Lago tion of the variety of squamation patterns Pequefno and Bahia de Puno, two smaller, exhibited by various species groups, and their shallower (to 40 m) basins. Most species are usefulness in formulating a hypothesis of the known only from the two smaller basins, but phylogenetic history of the genus. whether that represents the true distribution Unfortunately most collections prior to the of Orestias or merely the ease of collection, Percy Sladen Expedition have incomplete or is unknown. vague locality data. Thus, the majority ofma- In an effort to increase the productivity of terial known from outside the Titicaca Basin Lago Titicaca, introductions of brown trout cannot be used to describe precise ranges. (Salmo trutta fario) in 1939, and rainbow Since the diagnosis of Orestias by Valen- trout (Salmo gairdnerl) in 1941 were begun ciennes (1839), its monophyly has never been 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 113 challenged. The genus was defined by Parenti (Hennig, 1966) I formulate a hypothesis for (1981) on the following three synapomorphic the phylogeny of the 43 currently recognized characters: pelvic fins and girdles absent; species (figs. 26-31). Since the revision of Or- moderately lowset pectoral girdle lacks a first estias by Tchernavin (1944a), the Titicaca postcleithrum; and, vomer absent. Four ad- assemblage has been referred to or treated as ditional, newly discovered or described de- a so-called species flock (e.g., Villwock, 1962; fining characters are: cartilaginous middle Kosswig and Villwock, 1964; Richerson, dorsal and middle anal fin radials; ventral Widmer, and Kittel, 1977). The Orestias separation of the posterior and anterior cer- species in Lago Titicaca or the Titicaca Basin atohyals by a large gap filled with cartilage; do not form a monophyletic group, however. a unique squamation and head pore pattern; Hence, the concept of Titicaca supporting a and, the absence of a ventral extension on flock comprising all Orestias is refuted if the anguloarticular. A single median ovary monophyly is considered an essential part of or testis was mentioned by Valenciennes the definition of a flock. Only a few recent (1839) and subsequent workers as possibly discussions have questioned the application diagnostic of Orestias. A comparison of the of the species flock concept to any group of gonad morphology among Orestias and the related organisms in a lake (e.g., Reid, 1980, genera of Anatolian cyprinodonts, some of on Lake Lanao cyprinids) or treated the con- which also share this character, is made in cept of a flock as a useful way of referring to the present study. a group of species in a lake prior to the ac- In a recent revision of the killifishes, the ceptance or rejection of monophyly (e.g., order Cyprinodontiformes (Parenti, 1981), I Greenwood, 1979, and Stiassny, 1981, on Af- suggested that Orestias has no close South rican Rift Lake cichlids). American relatives, but is rather closely re- Orestias species in Lago Titicaca might be lated to the Anatolian cyprinodonts, and most considered a flock ifthe definition ofthe term closely to Kosswigichthys (including Anato- species flock includes a great diversity ofform lichthys). Apart from Eigenmann's suggestion and habitat. I defer a discussion ofthe species (1920) that Orestias might be closely related flock concept and its relationship to the phy- to the North American Empetrichthys (placed logeny and biogeography of Orestias species in the family Goodeidae by Parenti, 1981), to another paper (Parenti, in press). and that of Foster (1967) that Orestias could Thus, the aims of the present paper are to be closely related to the South American review the evidence for the relationship of aplocheiloid Cynolebias, this represented the Orestias to the Anatolian cyprinodonts and only detailed statement concerning the rela- the rest of the subfamily Cyprinodontinae; tionship of Orestias to other killifishes.2 In describe in detail characters (such as those of the present paper, I review briefly the rela- squamation and osteology) that are useful to tionship of Orestias to the Anatolian cypri- hypothesize phylogenetic relationships among nodonts, and the relationship of this larger the species, but have not been used exten- group to the North, Middle, and South Amer- sively in previous studies on the genus; di- ican cyprinodontines, which together with the agnose the recognizable species in addition Cuban and Jamaican Cubanichthys, com- to describing the new species; and present a prise the family Cyprinodontidae (sensu Par- detailed phylogenetic analysis of the species, enti, 1981, see table 2). as well as a key to their identification. Using the methods ofphylogenetic analysis ACKNOWLEDGMENTS The impetus for thus study was provided 2Eigenmann (1927, p. 52), without comment, de- by Dr. Peter J. Richerson and Mr. Tom Coon, scribed the range of Orestias as "Death Valley ofNevada and of the high plateau of Peru, Bolivia and Chile," and of the University of California, Davis, who therefore, without presenting a formal synonymy, con- sent a small sample ofOrestias collected from sidered Empetrichthys ofNevada to be ajunior synonym Lago Titicaca in the mid-1970s to the De- of Orestias. Most workers, however, followed Myers's partment ofIchthyology, American Museum 1931 suggestion that the two genera are not closely re- of Natural History with the promise of col- lated. lecting additional specimens while carrying 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

out a limnological study ofthe lake. In 1979, visits, I thank Dr. Donn E. Rosen, Ms. M. Mr. Coon expertly accumulated nearly 4000 Norma Feinberg, and Ms. Carol Schleifer of specimens of Orestias, Trichomycterus and the American Museum of Natural History, the introduced Basilichthys, from the Titi- Dr. William Smith-Vaniz and Mr. William caca Basin. I am grateful for the support of Saul ofthe Academy ofNatural Sciences, Drs. Dr. Richerson and Mr. Coon, and also that Keith Banister, P. Humphry Greenwood, Ms. of Sr. L. Edgar Farfan Vizcardo, director of Bernice Brewster, Mr. James Chambers and the Laboratorio de Puno, Instituto del Mar Mr. Gordon Howes, of the British Museum del Peru. They have all helped to inspire a (Natural History), Dr. William N. Esch- cooperative study of the Titicaca Basin. meyer, Mr. Michael Hearne and Ms. Pearl I am equally grateful to Mr. Tim Hardin, Sonoda, of the California Academy of Sci- of Colorado State University, for collecting ences, Drs. Robert K. Johnson, Donald J. and donating Orestias specimens from north- Stewart and Mr. Christopher Spurrier, of the ern Peru. Field Museum of Natural History, Dr. Wil- The taxonomic revision was initiated dur- liam L. Fink and Mr. Karsten Hartel, of the ing the tenure of a Smithsonian Postdoctoral Museum of Comparative Zoology, Dr. Ma- Fellowship at the National Museum of Nat- rie-Louise Bauchot and Ms. Martine De- ural History, Smithsonian Institution, and soutter, of the Museum National d'Histoire completed during the tenure of a NATO Naturelle, Dr. Martien van Oijen, of the Postdoctoral Fellowship at the British Mu- Rijksmuseum van Natuurlijke Historie, Dr. seum (Natural History). I thank my scientific Robert R. Miller and Ms. Alexandra Snyder- advisers, Drs. Stanley H. Weitzman and Creighton of the Museum of Zoology, Uni- Richard P. Vari (USNM) and Dr. P. Hum- versity of Michigan, and Drs. Stanley H. phry Greenwood (BMNH) for their encour- Weitzman, Richard P. Vari, and Ms. Susan agement, advice, support, and discussions. Jewett of the National Museum of Natural This project has also benefited from the ex- History, Smithsonian Institution. pert technical assistance and advice given un- Mr. Richard Mayden and Mr. John Sim- stintingly by colleagues at several institu- mons of the University of Kansas, Museum tions, including Ms. Janet Gomon and Ms. of Natural History, prepared figures 52 and Susan Jewett (USNM), Ms. Bernice Brewster, 53. These were supplied through the courtesy Mr. James Chambers, Mr. Phil Crabbe, and and cooperation ofDrs. Arratia-Schultze and Mr. Gordon Howes (BMNH), Ms. J. Abel E. 0. Wiley of the University of Kansas. (MNHN), and Mr. Michael Hearne and Ms. Much of the manuscript was patiently and Pearl M. Sonoda (CAS). professionally typed by Ms. Noreen Brews- I thank Drs. Gloria Arratia-Schultze and ter. Laurent Lauzanne for their willingness to ex- The manuscript benefited greatly from the change information and freely discuss their comments of several reviewers. work on Orestias. For reading and commenting on sections ABBREVIATIONS ofthe typescript, I thank Drs. Brian Coad, P. INSTITUTIONAL H. Greenwood, Mr. Gordon Howes, and Dr. AMNH, American Museum of Natural History, Ethelwynn Trewavas. Dr. Trewavas pains- New York takingly reviewed and edited the species ANSP, Academy of Natural Sciences, Philadel- etymologies and was always willing to discuss phia any and all aspects of the project. I also ben- BMNH, British Museum (Natural History), Lon- efited from her information on the work of don Dr. V. V. Tchernavin, who she saw carry out CAS, California Academy of Sciences, San Fran- a revision of Orestias 40 years ago at the cisco FMNH, Field Museum of Natural History, Chi- British Museum (Natural History). I must cago underscore my thanks to her for a host of IU, Indiana University (now at CAS, FMNH, kindnesses. UMMZ, and USNM) For information and the loan ofspecimens, KU, University of Kansas, Museum of Natural and in certain cases accommodation during History, Lawrence 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 115

MCZ, Museum of Comparative Zoology, Cam- Hb 1-3, hypobranchial 1-3 bridge Hs, hemal spines MNHN, Museum National d'Histoire Naturelle, lac, interarcual cartilage Paris Ih, interhyal RMNH, Rijksmuseum van Natuurlijke Historie, Le, lateral ethmoid Leiden Max, maxilla SU, Stanford University (now at CAS) Mes, mesethmoid UMMZ, University of Michigan, Museum of Zo- Mr 2, middle anal radial 2 ology, Ann Arbor Pas, parasphenoid USNM, National Museum of Natural History, Pb 1-3, pharyngobranchial 1-3 Smithsonian Institution, Washington, D.C. Pc, posterior ceratohyal Pcl, postcleithrum ANATOMICAL Pmax, premaxilla Aa, anguloarticular Pr 1, proximal anal radial 1 Ac, anterior ceratohyal Pro, prootic Afr, anal fin rays Pte, pterotic Apal, autopalatine Ptt, posttemporal B 1, 2, basibranchial 1, 2 Qua, quadrate Boc, basioccipital Rad, radials Br, branchiostegal rays Ret, retroarticular Cb 1-5, ceratobranchial 1-5 Sca, scapula Cl, cleithrum Scl, supracleithrum Cor, coracoid Sph, sphenotic Den, dentary Tpb 3, 4, toothplates ofthird and fourth pharyngo- E 1-4, epibranchial 1-4 branchials End, endopterygoid Vhh, ventral hypohyal Exo, exoccipital Open circles: cartilage Fro, frontal Stippling: bone

NOTE ON MATERIALS AND METHODS To carry out the aims of this study, it was Parenti (1981) for atherinomorph fishes in necessary to adopt a method of phylogeny general. Ontogenetic series were also exam- reconstruction with the goal of creating a ge- ined when available, for determination of nealogy of Orestias species that best repre- character state polarity. sents the one underlying, natural hierarchy. Methods of counts and measurements are Such a method is necessary to test hypotheses those proposed by Miller (1948) and Parenti concerning species flocks and patterns ofdis- (1981). All measurements are of straight-line tribution, as well as hypotheses of character distances recorded with metric dial calipers. evolution among cyprinodontid fishes. The I acknowledge the cautions of many workers method ofphylogeny reconstruction adopted on the use of ratios as they have been pre- here is that of phylogenetic systematics or sented traditionally in systematic studies (see cladistics (formally presented by Hennig, Humphries et al., 1981 for a brief review). 1966 and modified by many subsequent Tchernavin (1944a) presented a series of ra- workers). The cladistic methodology is con- tios based on several measurements, taken sidered the best currently available for pos- from a lateral view, as characters to distin- tulating such genealogical relationships (see guish among taxa. I do not use many of the Wiley, 1981 for a summary of the rationale descriptive characters ofTchernavin, not only and methods of phylogenetic systematics). because they represent ratios, but because they Outgroup comparisons for determining were not useful for discriminating among the character polarity in Orestias are made by species. This is not to say that measurements referring to related killifish genera, as well as cannot be used to discriminate among taxa. to characters described in Parenti (1981) for Measurements are important when no other cyprinodontiform fishes, and in Rosen and data, or only ambiguous data, exist. 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

Measurements were used primarily to aid tatives of many species were prepared ac- in identification ofspecies in conjunction with cording to the alcian blue-alizarin Red S descriptions of what I defined as consistent method outlined by Dingerkus and Uhler differences, such as, for example, wide head (1977). All descriptions of cartilaginous (as in Orestias luteus, fig. 7) versus a narrow structures are based on the examination of head. such preparations with comparisons made Description of osteological states follows with the collections of solely alizarin Red Parenti (1981) except as noted. S-stained preparations of cyprinodontiforms Counterstained preparations of represen- in the BMNH and USNM.

RELATIONSHIPS OF KILLIFISHES OF THE TRIBE ORESTIINI The genus Orestias, along with the Ana- The inferred most primitive state of the tolian cyprinodonts distributed in fresh and fifth character, tricuspid uniserial outer teeth, brackish waters throughout the Mediterra- is found among all Cyprinodontini and with- nean and Anatolia (see Parenti, 1981, p. 458, in the Orestiini, except for one species of fig. 86, for distribution map), comprise the Kosswigichthys (K. asquamatus) and Ores- tribe Orestiini. Together with the North, tias. In Orestias, the outer teeth have been Middle, and northern South American cy- described as unicuspid or bicuspid and are prinodonts of the tribe Cyprinodontini, they in an irregular row, which has been inter- comprise the subfamily Cyprinodontinae, preted as two or more rows in particular which, with its sister group, the subfamily species. Cubanichthyinae, containing the genus Cu- Franz and Villwock (1972) studied tooth banichthys of Cuba and Jamaica, comprise development in Anatolian cyprinodonts, and the family Cyprinodontidae sensu stricto (ta- recognized three developmental stages: a sin- ble 1). gle row of unicuspid teeth followed in de- Relationships among the genera ofthe Or- velopment by an anterior, single row of tri- estiini, and their relationship to other cy- cuspid teeth, and finally by the addition of a prinodontids are presented in figure 2; the labial row of tricuspid teeth. Typically in character numbers in that figure correspond Aphanius, the most interior of the unicuspid with those of the following discussion. teeth are lost, the middle row of tricuspid Fishes of the subfamily Cyrpinodontinae teeth persists as the prominent adult denti- form a monophyletic group defined by the tion, and the outer row is what is generally following five synapomorphic characters referred to as the replacement dentition. In (Parenti, 1981, p. 521): (1) the second pha- Kosswigichthys asquamatus, the two rows ryngobranchial is offset relative to the third; forming anterior to the first in ontogeny con- (2) Meckel's cartilage is expanded posterior- sist of unicuspid rather than tricuspid teeth, ly; (3) the transverse processes of the verte- and the interior, as well as the two anterior brae are reduced and cup-shaped; (4) pari- rows, persists in adults. Villwock (1976) de- etals are absent; and (5) outer jaw teeth scribes the irregular placement ofteeth in the generally are found in a single row and prim- upper and lower jaws of Kosswigichthys as a itively are tricuspid. "slot-link" formation. Character number 3, reduced and cup- No ontogenetic studies are known to have shaped transverse processes ofthe vertebrae, been published on outer tooth formation occurs in many but not all species of Orestias within the Cyprinodontini. Adult members (e.g., they are small in Orestias hardini). The of this tribe typically have a single row of somewhat elongate transverse processes in tricuspid outer jaw teeth, with several tricus- some Orestias are therefore considered to be pid labial replacement teeth. A juvenile Cy- secondarily derived. prinodon variegatus (BMNH 1948.8.6:1634- 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 117

TABLE 1 The Family Cyprinodontidae I z C.) P- (sensu Parenti, 1981) I z 0 Cn C') a Family Cyprinodontidae C,) 0 z z Subfamily Cubanichthyinae cc 4 z escf) C, Genus Cubanichthys w IL a.I a. Subfamily Cyprinodontinae Tribe Orestiini 27 Genus Aphanius 26 25 Genus "Aphanius" 24 Genus Kosswigichthys 23 Genus Orestias 22 Tribe Cyprinodontini 0 ---a21z Genus Cyprinodon Li& b rl20 Genus Jordanella L I t-_-019 Genus Cualac 0- 0 18 Genus Floridichthys 0| ? 17 Genus Megupsilon P-Ib-r -r aI1a1 II I T O0 14 0 - ° - ;13 I 0 0- r1 1665, 18.8 mm SL) examined has a similar dental arrangement; there is no row of inte- u rior jaw teeth. On the basis of ontogenetic patterns alone, one could describe the transition of cyprino- u , dontine outer jaw dentition as from the presence of two rows and an outer replacement I a 6 row to a loss of the most interior row. How- 5 ever, on the basis of the defined monophyly 4 ofthe subfamily Cyprinodontinae, and on the 3 distribution of all other known derived char- 2 acters within the subfamily, I interpret the 1 state of a single row of tricuspid teeth with an outer replacement row to be derived at FIG. 2. Synapomorphy diagram of groups the subfamily level, and the unicuspid, multi- within the subfamily Cyprinodontinae (sensu Pa- row adult teeth of Kosswigichthys and Or- renti, 1981). Characters are numbered as those in estias to be a secondarily derived state. Usu- the text. Black square represents synapomorphy, ally, such a character would be termed a "re- open square symplesiomorphy, and a half-square versal" to a more primitive state. However, synapomorphy found injust some members ofthe because ofwhat is known about the ontogeny terminal taxon (see text for discussion). of tooth formation in Anatolian cyprinodonts, and the pattern ofdentition in the tribe Cyprinodontini, it is most parsimonious to development (e.g., Ermin, 1946; Villwock, describe the derived state in Kosswigichthys 1976; Grimm, 1979). and Orestias as the absence of the suppres- Villwock (1976, p. 23) summarized the sion of the most interior row. phenomenon of reductions in Anatolian cy- Reduction or loss of characters is a com- prinodonts: "Populations living under more mon phenomenon among killifishes in gen- or less normal fresh-water conditions look eral, and among the Orestiini in particular in like any Aphanius from other Anatolian fresh- which there are often reductions in squa- water systems. They show normal scaliness, mation, tooth cusp number and pelvic fin normally shaped tricuspid teeth and no fin 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 reductions. However, those Aphanius which hyal was not given as diagnostic of "Aphani- live in the lake waters containing sulfate, show us" (Parenti, 1981, p. 523), its presence has a high degree of scale-reduction, alterations been confirmed in additional material of"A." in fin rays, or even total absence ofpelvic fins mento (BMNH 1920.3.3:203-222). and partial reduction in the lateral tips of In primitive species ofthe genus Aphanius their tricuspid teeth." and in the species of the Cyprinodontini the Characters such as reduction in squama- head pore and squamation pattern, consid- tion, or absence of pelvic fins are assessed as ered here to be the primitive pattern of the derived at those levels where they are invar- Cyprinodontinae, is characterized by a se- iant. For example, no species of Orestias has ries of pores and scales arranged in a partic- pelvic fins or fin girdles at any developmental ular pattern that varies somewhat among stage. I use it as a derived character because species (fig. 3A, B, C). The patterns of pre- it is invariant within the genus and also be- opercular and mandibular pores, are also rel- cause pelvic fins are found in the hypothe- atively invariant. In the derived pattern, seen sized closest relative, Kosswigichthys. in its most primitive state in "A." mento (fig. The tribe Orestiini is defined as monophy- 3D, E, F), there are no distinct pores; instead, letic by one synapomorphic character: (6) a there are minute neuromasts dorsally which broadly expanded medial process ofthe den- also continue along the preopercle and lower tary (Parenti, 1981, p. 413, fig. 43) which jaw. gives the orestiines a characteristically robust Kosswigichthys asquamatus is unscaled; the lower jaw. other species in the genus exhibit the derived Anatolian cyprinodonts have often all been state of the pore and squamation pattern as placed in the genus Aphanius (e.g., Villwock, in "A." mento. 1976); there are over 30 nominal species, of The head pore pattern exhibits little vari- which nine are recognized as valid by Vill- ation among Orestias species. General char- wock. I was unable to define the Anatolian acteristics of the pattern include a series of cyprinodonts as a monophyletic group, and minute neuromasts arranged in a "lyre-shape" pointed out four derived characters that some because it suggests the outline of a lyre. The species of Aphanius and the genus Kosswig- dorsal cephalic neuromasts (fig. 3G) are in ichthys (including Anatolichthys) share with one or two rows across the snout and contin- Orestias (Parenti, 1981, p. 457): (7) a cartilag- ue posteriorly, medial to the posterior nares, inous interhyal; (8) a unique head sensory along the dorsal surface ofthe head lateral to pore pattern represented by neuromasts; (9) the dorsal head scales. The neuromast series a reduced or absent dermosphenotic; and (10) continues medially, then laterally along the an upturned lower jaw associated with the preopercle and mandible. There are no preor- embedding ofthe protractor hyoidei between bital pores; a series of neuromasts encircles the jaw suspensoria. An additional synapo- the eye (fig. 3H). A similar neuromast pattern morphic character that I discuss is: (11) ova- occurs in the South American aplocheiloid ries and testes single in adults. Cynolebias (Parenti, 1981, p. 371, fig. 13G, The species ofAphanius that were assessed H, I) that has been assessed as convergent as being more closely related to Kosswigich- because Cynolebias clearly has the derived thys and Orestias than to the rest of the characters of the suborder Aplocheiloidei, Aphanius species could not be defined as family Rivulidae; whereas, Orestias has the monophyletic. However, in order to empha- derived characters of the suborder Cyprino- size their close relationship to the Kosswig- dontoidei, family Cyprinodontidae. ichthys-Orestias sister group pair, they were In Aphanius fasciatus as well as in the referred to as "Aphanius" (Parenti, 1981, p. species of the Cyprinodontini, the dermo- 522). I considered Aphanius mento to be rep- sphenotic is a long bone with a deep trough resentative of this group; other species of for the sensory canal. In genera that exhibit Aphanius that are more closely related to the derived neuromast pattern, the dermo- Kosswigichthys and Orestias include chantrei sphenotic is very reduced (e.g., as in "Aphani- and sophiae. Although a cartilaginous inter- us" and some Orestias) or absent (e.g., as in 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 119

FIG. 3. Aphanius fasciatus (BMNH 1951.2.6:1-37): A. lateral view of preopercular and preorbital pores, B. dorsal view of supraorbital pores and squamation, C. ventral view of mandibular and preopercular pores; "Aphanius" mento (BMNH 1968.12.13:431-435): D. lateral view ofneuromast pattern, E. dorsal view of neuromast pattern and squamation, F. ventral view of neuromast pattern; Orestias cuvieri (BMNH 1944.6.6:1-6): G. lateral view ofneuromast pattern, H. dorsal view ofneuromast pattern and squamation, I. ventral view ofneuromast pattern. Diagrammatic representations ofcephalic sensory pore and squamation patterns. 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

ribs

A prl

mr 2

afr

prl1

afr

FIG. 4. A. Orestias agassii (AMNH 52147); B. Orestias tomcooni (AMNH 52189). Diagrammatic representation of internal anal fin supports, anterior is to the left. some Kosswigichthys and other Orestias). and morphology of the gonads of Anatolian When present in these genera, it has no trough cyprinodonts. Concerning the development and therefore carries no sensory canal. ofthe gonads in Aphaniusfasciatus, he stated Species of"Aphanius," Kosswigichthys, and (pp. 180-18 1): "The ovary consists of two Orestias have an upturned lower jaw that, in components, the right and left gonad. Even lateral view, in some cases is perpendicular the gonad of almost mature females retains to the body axis. A character correlated with its original paired form very clearly .... The this upturned lower jaw is the protractor development of the testes proceeds much hyoidei embedded between the jaw suspen- slower than that of the ovaries and they are soria. This character was described by Paren- usually ofsmaller size. Although the right and ti (1981, p. 457) as an embedded urohyal, but left testes of the male gonad are juxtaposed, is considered here to be more properly de- they are normally not close to each other as scribed as a muscle, rather than a bone, char- the testes of Aphanius sophiae." Concerning acter. the development of the gonads of Aphanius Valenciennes (1839) mentioned the united cypris (=mento), Konuralp stated (p. 179): ovaries and united testes as a character of "The left and right ovaries are united in the Orestias. Tchernavin (1 944a, p. 149) includ- female .... The primary ovarian cavities of ed in a series ofcharacters for Orestias "ovary the right and left side join each other in order and testis single." An examination has con- to form a common secondary ovarian cavity firmed the presence of this character in Or- .... In the male, too, there are two gonads, estias. one on the right and one on the left. These Konuralp (1955) studied the development gonads can be distinguished from one another 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 121

27 vertebrae). However, a specimen of Aphanius fasciatus, BMNH 1951.2.6:1-37, has 28. Also vertebral counts among the Cy- prinodontini range from 24 to 29 (personal observ., Miller, 1956; Miller and Walters, 1972). The consistently high number of vertebrae in Orestias (30 or more) may be an additional defining character of the genus; however, our present poor knowledge of the relationships of species of Anatolian cypri- , ...\. nodonts makes the determination ofpolarity FIG. 5. Cyprinodon variegatus (BMNH of vertebral counts problematic. 1948.8.6:1634-1665). Diagrammatic representa- As I said before, a character such as 13 tion of hyoid bar. V above, "reduction or absence of scales" is used only when it is consistent among members of the considered taxa. Particular pat- but they are united." Konuralp also surveyed terns ofscale distribution are used in the Phy- gonad morphology and development in logenetic Analysis section to define groups of Kosswigichthys and Anatolichthys and con- Orestias species. cluded that the gonads were united as in "A." Three characters (19-21) are in conflict with mento. [Anatolichthys, however, has some the scheme of relationships as depicted in unique characteristics ofgonad morphology.] figure 2. In all Anatolian cyprinodonts United gonads occur in other cyprinodontiform fishes, e.g., in viviparous goodeids (Hubbs and Turner, 1939). However, Konu- ralp (1955) considered the formation of the gonads of Anatolian cyprinodonts to be unique among cyprinodontids (equals oviparous cyprinodontiforms). Because goodeids are distantly related to orestiines, the early union ofthe gonads in Orestias, "Aphanius," and Kosswigichthys is treated most parsimoniously as a synapomorphy of the three genera. Kosswigichthys shares two derived characters with Orestias: (12) the posttemporal bone is straight, that is, the ventral limb is represented by a ligament; and (13) there is VI a reduction or absence of scales. A third character, an increase in the number of vertebrae to 28 or more as opposed to a range of 24 to 27 among other members of the Cyprinodontinae, was used by Parenti (1981, p. 458) to unite Kosswigichthys and Orestias into a monophyletic group. Further B investigation has shown that the level at which an increase in the number of vertebrae can be used as a derived character is ambiguous. BR Representatives ofoutgroups have lower vertebral counts (e.g., "Aphanius" mento, BMNH FIG. 6. A. Orestias pentlandii (SU 9306); B. 1920. 3.3:203-222, has 25 vertebrae; Apha- Orestias luteus (AMNH 52130). Diagrammatic nius dispar, BMNH 1977.12.13:1-490, has representation of hyoid bar. 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

(Aphanius, "Aphanius," and Kosswigich- Therefore, the genus Orestias may be de- thys), the mesethmoid is cartilaginous (char- fined by seven synapomorphies: (21) pelvic acter 19) and never ossified as it is in most fins and fin girdles are absent; (22) the pec- other cyprinodontiforms, including Orestias toral girdle lacks a first postcleithrum; (23) and the Cyprinodontini. Eight characters the vomer is absent; (24) the middle dorsal (seven through 14) oppose the formation of and middle anal radials are cartilaginous (fig. a group based on this single character. 4); (25) the anterior and posterior ceratohyal In all Orestias species and in some species are separated ventrally by a large gap of car- of the Cyprinodontini, the third and fourth tilage (fig. 6) as opposed to the usual cyprino- pharyngobranchial toothplates are fused to dontiform condition in which they are close form one large tooth-bearing element of the together (fig. 5); (26) there is a unique squa- dorsal gill arches (character 20). The distri- mation and head pore pattern, described in bution ofthis character, and also the fact that the Phylogenetic Analysis, and (27) the an- it is found among other groups of cyprino- gularticular lacks a ventral extension parallel dontiforms, makes it of questionable value to the retroarticular (see Parenti, 198 1, p. 41 3; in determining relationships. At this stage in fig. 43). our knowledge of cyprinodontiform system- Pelvic fins and fin girdles are absent in some atics, it certainly should not be used to hy- members ofthe Cyprinodontini and Aphani- pothesize that some species of the Cyprino- us species, as well as in all Orestias (character dontini are more closely related to Orestias. 21). The absence is used nevertheless as a Also, the tribe Cyprinodontini is a well-de- defining character of Orestias because its pre- fined monophyletic group. Its defining char- sumed closest relative, Kosswigichthys, has acters are: (15) the supraoccipital forms the pelvic fins and fin girdles under normal cir- dorsal wall ofthe foramen magnum; (16) the cumstances. Also, the character is corrobo- neurapophyses ofthe first vertebra are angled rated by the existence of six additional de- anteriorly and firmly applied to the skull; (17) rived characters (22 through 27). the exoccipital condyles are absent; and (18) the pharyngobranchial teeth are distributed in discrete rows.

PHYLOGENETIC ANALYSIS In the last and most comprehensive treated 0. mulleri and crawfordi, the fully scaled, ment ofOrestias by Tchernavin (1 944a), there relatively deepwater species ofLago Titicaca. was no formal statement of relationships Relationships among the four groups were among all species. Tchernavin divided the not specified, although Tchernavin's key and species into four groups, referred to as Groups formal text implied that the incompletely I to IV, each distinguished by overall body scaled species of Groups I to III were a group shape or squamation pattern. Group I of distinct from the fully scaled species ofGroup Tchernavin comprised 0. cuvieri and pent- IV. Also, a precise definition using characters landii, the large, elongate midwater species that today we would recognize as synapom- of Lago Titicaca. Group II included species ophies for each group was not presented. In such as 0. agassii and luteus which were spite of this, Tchernavin's groupings repre- characterized by relatively deep caudal pe- sent a reasonable first approximation of duncles and by the anterior ribs becoming monophyletic groups within the genus, the large and swollen with age (i.e., increase in definition and composition of which are Standard Length) to form very wide-bodied specified more rigorously in the present re- adults in some species (see fig. 7). Group III vision. included 0. gilsoni and tutini, relatively small, The major monophyletic groups recog- inshore species of Lago Titicaca character- nized are referred to as species complexes, ized by large eyes that project dorsally above the composition of which are listed in table the primary dorsal profile. Group IV includ- 2. Group I of Tchernavin has been found to 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 123 be monophyletic and is referred to as the cu- TABLE 2 vieri complex. Two species described recently Orestias species Recognized in the by Lauzanne (198 1), 0. ispi and 0. forgeti, Present Study, Grouped into Species Complexes are included. cuvieri complex agassii complex Groups III and IV together form a well- cuvieri agassii defined monophyletic group, although nei- pentlandi empyraeus ther of Tchernavin's groups is definable as ispi frontosus monophyletic. Two monophyletic groups of forgeti polonorum species are recognized here, and referred to elegans as the gilsoni and mulleri complexes. mulleri complex jussiei Group II is referred to as the agassii com- gracilis puni plex. One ofthe group's subgroups comprises mulleri parinacotensis five species that share a derived squamation crawfordi laucaensis tutini tschudii pattern. Because in the following discussion gymnotus the pattern is compared with that of other incae hardini species in the agassii complex as well as other ctenolepis species complexes, the five species together gilsoni complex ascotanensis (luteus, rotundipinnis, farfani, albus, and oli- gilsoni richersoni vaceus) are referred to as the luteus group. taquiri multiporis Allen (Eigenmann and Allen, 1942) pre- mooni mundus sented a dendrogram of Orestias species along uruni ututo with his succinct the genus. Be- minimus silustani revision of minutus luteus cause the relationships were based primarily tchernavini rotundipinnis on overall phenetic similarity, and because tomcooni farfani there are several obvious misidentifications imarpe albus of species resulting in an unsupportable robustus olivaceus scheme ofrelationships, the information con- tained in the dendrogram is unusable for phylogenetic inference. Therefore, it is not ana- lyzed or further commented on here. the defining characters of Orestias is its unique Selected characters or character complexes squamation and head pore pattern. A prom- used to define species or groups of species are inent feature of the pattern is a distinct me- explained below; additional characters used dian dorsal ridge of scales that covers the to formulate a hypothesis ofthe relationships middorsal surface of the body from approx- of Orestias species are discussed in the syn- imately the posterior limit ofthe head to the apomorphy scheme (pp. 150-160), and in the anterior base of the dorsal fin (see fig. 7). diagnosis of each species. When a character These scales are generally larger and thicker is invariant within a complex, it is described than surrounding scales, and are almost al- in the following discussion as being present ways arranged in a straight, median line. Cor- in the complex rather than being described respondingly, scales on either side ofthe dor- for each species. Similarly, when a character sal ridge are relatively thin, or may be absent is primitive for a complex, it is described for altogether. The latter condition results in na- the complex with remarks about its distri- ked areas between the ridge and the lateral bution within Orestias. body scales in adults of distantly related SQUAMATION3 AND NEUROMAST PATTERN: species in the cuvieri and agassii complexes, As briefly discussed in the section on rela- although individual variation within species tionships within the tribe Orestiini, one of is marked. Variations on the general pattern ofa well-formed dorsal ridge occur in several groups of species, as detailed below and in 3In the present paper, the term squamation refers to the synapomorphy scheme. types and patterns of distribution of scales on the body. The dorsal surface of the head is incom- The term scalation is considered to be a synonym of pletely covered with irregularly distributed squamation and therefore is not used here. thickened scales that have been referred to 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 7. Orestias luteus (AMNH 52134), 110.0 mm SL. Lateral, dorsal, and ventral views, from top to bottom, of an adult female.

as scutes in Orestias (Tchernavin, 1 944a, granulations are always present in subadults 1946) although the term scutes is considered and adults of the luteus group of the agassli inappropriate here. These head scales may be complex, that is, in luteus, rotundipinnis, far- granulated, as they often are in adult cuvieri fani, albus, and olivaceus. and pentlandii of the cuvieri complex. Such The presence of thickened and granulated 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 125 scales might be considered a synapomorphic character of the entire genus that has been lost or modified in the gilsoni and mulleri complexes, ispi andforgeti ofthe cuvieri complex, and the agassii complex minus the species of the luteus group. However, it is most parsimoniously treated as independently derived within the luteus group and in cuvieri andpentlandii. Along with ispi, these last two species form a monophyletic subgroup of the cuvieri complex with a defining character of very numerous, small lateral scales (see discussion of meristic characters, below). Another modification ofhead scales occurs in a group of species within the agassii complex. Ten species (jussiei, puni, tschudii, frontosus, ctenolepis, richersoni, multiporis, polonorum, empyraeus, and agassii) are distinguished readily by having all scales on the head and the dorsal and lateral surfaces of the body anterior to the dorsal fin ex- C D tremely smooth and barely overlapping. The dorsal scales are the thickest, are smooth, and FIG. 8. Outline of anterior view of head A. Orestias agassii, B. Orestias puni, C. Orestias al- have few or no discernible concentric striae. bus, and D. Orestias luteus, to illustrate differences The five species ofthe luteus group are further in shape of head in wide-headed species. distinguished from the rest of the agassii complex species by having granulated scales. The derived neuromast pattern of Orestias to the fact that the totally unscaled Kosswig- species (fig. 3G, H, I) described in the section ichthys asquamatus is a member of the sister on relationships within the tribe Orestiini is genus of Orestias. However, fully scaled very nearly obscured in large adults of these species are not scaled in the same manner as species by the abutting edges ofthe irregularly other fully scaled cyprinodontines which have shaped scales. regular scale rows on the ventrum. Fully In all species of the agassii complex, the scaled Orestias have very small, thin, and body scales immediately posterior to the pec- irregularly distributed scales on the ventrum toral fin base are relatively large and form and on the basal portion ofthe pectoral, dor- what will be referred to as a lateral shield. In sal and anal fins; this I interpret as a derived the species of the luteus group that have a condition. Juveniles of fully scaled species very wide head, these scales form thickened, are fully or incompletely scaled, whereas ju- granulated shoulders (see figs. 7, 8). veniles of incompletely scaled species are in- In most species of Orestias, the ventral sur- completely and irregularly scaled. Thus, the face from the tip of the lower jaw to the base fully scaled condition is not primitive for the of the anal fin, as well as the basal portion of genus. the pectoral, dorsal, and anal fins are devoid Fully scaled species may be closely related of scales. A number of species of the mulleri to incompletely scaled species, as is the case complex, two species of the cuvieri complex with forgeti and ispi, which, along with pent- (ispi andforgeti), and 0. mooni ofthe gilsoni landii and cuvieri comprise the cuvieri com- complex, are fully scaled. A fully scaled con- plex. Together the last three species have a dition would initially be assessed as primitive synapomorphy of an increase in the number for Orestias because that is the primitive con- ofscales in a lateral series as well as a number dition for cyprinodontiform fishes. Problems ofother characters (see table 7 and discussion with interpretations of degree of squamation on meristic characters, below). The numer- have already been discussed with reference ous lateral body scales are thin and small. 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 10. Orestias pentlandii (SU 9306). Dia- grammatic representation of dorsal gill arches, ventral view.

TPB3&4 FIG. 9. Orestias agassii (AMNH 52147). Dia- at the base of the caudal fin. (It should be grammatic representation of dorsal gill arches, noted that specimens of parinacotensis and ventral view. laucaensis were not examined in this study; information is from Arratia, 1982, and the photographs of figs. 52 and 53). Among Orestias species, ctenii are often In general, larger species tend to be irreg- present on scales of males, but are usually ularly scaled, whereas smaller species tend to irregularly distributed and weakly formed. be fully scaled. Like all generalizations, how- One species of the agassii complex from the ever, this is not always true, and therefore Titicaca Basin (0. ctenolepis) is defined by the phylogenetic significance of incomplete having numerous ctenii on all head scales and squamation of Orestias remains problematic. all lateral body scales in males (fig. 59), ex- Unique patterns which include incomplete cluding those of the lateral shield. As many squamation can be described, however. as 10 ctenii per scale have been observed on Five species within the agassii complex lateral body scales. (gymnotus, ascotanensis, elegans, parinaco- HYOBRANCHIAL APPARATUS: Gill arch tensis, and laucaensis) share a reduced squa- structure varies widely among Orestias mation pattern which has several unique species. General patterns ofgill arches as well characteristics. Aspects ofthe pattern may be as of the hyoid bar are described in this sec- seen best in the photograph of the holotype tion, with characteristics specific to species of 0. gymnotus (fig. 55). The dorsal surface or species groups detailed in the synapomor- of the head is irregularly scaled; the median phy scheme and systematic accounts. dorsal ridge is either fully present or repre- In all Orestias species, the third and fourth sented by just a few, randomly placed scales. pharyngobranchial toothplates are fused (figs. The ventral surface and basal portion of the 9-12). These toothplates are found fused in pectoral, dorsal, and anal fins are unscaled. other cyprinodontids (fig. 2, character 20), The lateral body scales are present in a unique and will not be used as a defining character pattern: beginning at the posttemporal, there of Orestias for reasons discussed above. are one or two uneven rows of lateral scales. The cuvieri complex comprises four species The number of scale rows increases poste- that are pelagic fishes in Lago Titicaca. They riorly so that there are four to five scale rows are plankton feeders (ispi, pentlandii, andfor- 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 127

FIG. 12. Orestias crawfordi (AMNH 52126). Diagrammatic representation ofdorsal gill arches, FIG. 11. Orestias crawfordi (AMNH 52126). Diagrammatic representation ofdorsal gill arches, dorsal view. ventral view. (fig. 9). In the mulleri and gilsoni complexes geti) or omnivores (cuvieri). In all four species, (fig. 11) the second pharyngobranchial is re- the ventral branchial apparatus is elongate duced to a U-shaped bone with a reduced (fig. 15). The basihyal bone is long and nar- toothplate that bears few or no teeth. Re- row, rather than short and triangular (fig. 13) duction in the number ofteeth in the anterior or rectangular (fig. 14) as it is in the Anatolian section of the dorsal gill arches is correlated and certain other cyprinodontiforms. The fifth with the formation of rather larger molari- ceratobranchials (lower pharyngeals) are small form teeth on the third and fourth pharyn- and narrow, possess numerous simple teeth, gobranchial toothplates in two species of the and are very close together or fused in adults. mulleri complex (0. crawfordi, fig. 11, and The dorsal gill arches are reduced in length 0. incae). In these species, the fifth cerato- relative to those of other species complexes branchials (lower pharyngeals) have molari- and other cyprinodontids. The gill rakers are form teeth as well (fig. 16), larger and blunter usually branched rather than simple, al- in crawfordi than in incae. though this character varies among individ- Both 0. crawfordi and incae are relatively uals. deepwater killifish (taken at depths to 38 m) A synapomorphic character of cyprino- that feed on molluscs. Several species in the dontoids described by Parenti (1981) is a re- luteus group (e.g., luteus, rotundipinnis, and duction of the interarcual cartilage to ap- possibly farfani) also have molariform teeth proximately one-half the length of the first on the fifth ceratobranchials and the third epibranchial. This is the case for Kosswigich- and fourth pharyngobranchial toothplates. thys asquamatus (BMNH 1948.3.15:40-43) The second pharyngobranchial toothplate in and most species of Orestias (fig. 9); however, these three species has a primitive dentition; in the cuvieri complex, the interarcual carti- that is, one or more rows of conical teeth are lage is increased in size to as long as, or longer present. than, the first epibranchial bone (fig. 10). The correlation of molariform pharyngeal In the primitive state, the second pharyn- teeth and fused fifth ceratobranchials with a gobranchial bears a toothplate which has a particular diet is not apparent among Ores- series of simple teeth in one or several rows tias species. Within the luteus group, 0. lu- 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 14. Orestias mulleri (AMNH 52118). Diagrammatic representation of ventral gill arch- FIG. 13. Orestias jussiei (CAS 46175). Dia- es, dorsal view. Ossified portion ofbasihyal black- grammatic representation of ventral gill arches, ened. Dotted line represents outline of fifth cera- dorsal view. Ossified portion of basihyal black- tobranchial toothplates. ened. Dotted line represents outline of fifth ceratobranchial toothplates. (fig. 12). Associated with this flange is a large adductor IV muscle that fills the cupped sur- teus has large pharyngeal teeth, many ofwhich face of the bone. It is inferred that the con- have rounded cusps, whereas the closely re- dition allows for more efficient crushing of lated 0. albus has blunt-tipped, unicuspid prey, although no functional analyses have pharyngeal teeth which could not properly be been carried out to test this hypothesis. referred to as molariform. Both species have The second epibranchial bone in some been reported to be molluscivores (Pellegrin, species of the cuvieri, mulleri, gilsoni, and 1904a), and 0. albus also to be a piscivore agassii complexes bears a distinct dorsally (Tchemavin, 1944a). directed process that is capped by a ball of Blunt pharyngeal teeth are apparently cor- cartilage (fig. 10). This process is different related with the crushing of prey; however, from the slight expansion ofthe second phar- fused fifth ceratobranchials (lower pharyn- yngobranchial bone that is considered to be geals) are not. Pellegrin (1 904a) reported fused primitive for acanthopterygian fishes (see lower pharyngeals in 0. /uteus, albus, tschu- Rosen and Parenti, 1981, fig. 1) because it is dii, and pentlandii. However, the lower pha- a distinct process that bears a cap ofcartilage. ryngeals are close together in the first three The process does not articulate with that of species, but fused only in the last, pentlandii, any other gill arch element, nor are the ad- and other species of the cuvieri complex. As ductor II or levator extemus II muscles mod- previously discussed, members of the cuvieri ified in any species examined that have the complex are either strictly planktivorous or process. The distribution of this character (it are omnivorous, indicating no correlation of is not present in all species of any complex) fusion of lower pharyngeals with diet. makes it problematic for use in a phyloge- An autapomorphy of 0. crawfordi is a large netic analysis; I do not use it here to define lateral flange on the fourth epibranchial bone any group of species. 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 129

FIG. 16. Orestias crawfordi (AMNH 52126). Diagrammatic representation of fifth ceratobranchials (lower pharnygeals), dorsal view.

makes their use in the phylogeny reconstruction questionable. I point them out here so that the distribution of such gill arch structures will become better known. Two species of the cuvieri complex (ispi CB5 and pentlandii) are defined as sister species FIG. 15. Orestias ispi (AMNH 52111). Dia- by an increase in the number and arrange- grammatic representation of ventral gill arches, ment of branchiostegal rays, among other dorsal view. Ossified portion of basihyal black- characters. In other Orestias, the Anatolian ened. Dotted line represents outline of fifth ceratobranchial toothplates. cyprinodonts and many other cyprinodontoids there are five branchiostegal rays. The primitive number of branchiostegal rays for In all but five species ofthe agassii complex cyprinodontiforms is six, with two hairlike examined (mundus, hardini, ututo, luteus, and rays present on the anterior arm of the an- albus), the first hypobranchial bone is divid- terior ceratohyal, separate from the four ed where it meets the first basibranchial (fig. bladelike rays on the posterior section of the 13). Both the anterior and posterior sections hyoid bar, as in Cyprinodon variegatus (fig. of the hypobranchial that meet the basi- 5). When the branchiostegal number is re- branchial have distinct cartilages. This con- duced to five among cyprinodontoids, it is dition is interpreted as derived because in due to a reduction in the number of anterior other cyprinodontids, including Kosswigich- hairlike rays (fig. 6B). In ispi and pentlandii thys, the first hypobranchial is simple, as it there is just one anterior ray; the number of is in Orestias mulleri (fig. 14). However, a branchiostegal rays is increased to six by the divided hypobranchial bone is found among addition of a fifth ray on the section of the some other cyprinodontoids as well. Thus, hyoid bar posterior to the notch (fig. 6A). the character of the divided first hypobran- FINS: Three synapomorphic character chial may be an additional derived character complexes of the fins have already been de- of the agassii complex that has secondarily scribed as defining Orestias. These are the attained a condition in the five species named absence ofpelvic fins and fin girdles; the pres- above similar to the primitive condition. ence ofcartilaginous rather than ossified mid- Both these gill arch characters (a process dle dorsal and anal fin radials (fig. 4); and the on the second epibranchial bone and a di- absence ofthe first postcleithral bone (fig. 17). vided first hypobranchial bone) are recog- A unique form ofthe structure and position nized as derived characters. However, the of the anal fin serves as a synapomorphic distribution of the characters within the ge- character of the mulleri and gilsoni com- nus Orestias and among other killifishes plexes. Typically among oviparous cyprino- 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

In the mulleri complex, the dorsal and anal fin base extend beyond the primary body profile, as in 0. crawfordi (fig. 38). In fishes of :1 this complex, the dorsal and anal inclinators are greatly enlarged. The gilsoni complex has one defining character, the caudal fin. Typically for killifishes, the caudal fin is rounded or truncate, and the procurrent caudal fin rays extend around the hypural plate in a nearly perfect semicircle Cl (fig. 18B). In the gilsoni complex, the procurrent caudal fin rays lie closer to the vertebral column giving the caudal fin what can be described as a tapered external outline (fig. 18A). Females of three species complexes (gilsoni, mulleri, and agassii) have a large anal pouch that extends over the base of the first several anal fin rays (e.g., figs. 7 and 69). There .1 is also a large, unpigmented anal papilla. The pouch and papilla are notably absent in 0. uruni of the gilsoni complex. FIG. 17. Orestias crawfordi (AMNH 52126). In three species ofthe cuvieri complex (cu- Diagrammatic representation of left pectoral gir- vieri, pentlandii, and ispi) the caudal fin is dle. Anterior is to the left. distinctly lunate, whereas in all other species of Orestias it is rounded or truncate. Killi- fishes generally have rounded or truncate dontiform fishes, the longitudinal axis of the caudal fins, although a lunate caudal fin is anal fin base is roughly parallel to the ver- found in, for example, the pelagic cyprino- tebral column. The proximal anal radials are dontoid of Lake Tanganyika, Lamprichthys therefore parallel to, and their tips extend tanganicanus (Boulenger). The lunate caudal between, the hemal spines (fig. 4A). The fin is interpreted as a derived character fur- proximal radials are all of approximately the ther indicating the closeness of relationship same length, becoming slightly shorter in the of the three cuvieri complex species named posterior section of the fin. above. In the mulleri and gilsoni complexes, the CHROMOSOMES: Chromosome numbers anal fin is modified such that the longitudinal have been reported in four species of Ores- axis ofthe base ofthe fin lies at an angle away tias. The primitive haploid number for cy- from the vertebral column (fig. 4B). The prinodontiforms is 24, and that is the number proximal anal radials do not lie between the reported for 0. luteus and 0. agassii (Lueken, hemal spines; they become relatively smaller 1962) and 0. parinacotensis (Arratia, 1982). anteroposteriorly. The fourth species, 0. laucaensis, reportedly A derived character for cyprinodontoids has a diploid chromosome number of 52 in (sensu Parenti, 1981) is large inclinators of males and 50 or 51 in females (Arratia, 1982). the anal fin that extend dorsally nearly to the Both the increase in, and the sexually di- division between the hypaxial and epaxial morphic nature of, chromosome number are musculature. This is the generalized condi- defining characters of this species. tion for Orestias species. In the mulleri and Sexual dimorphism of chromosome num- gilsoni complexes the anal inclinators are ber has been reported in other cyprinodon- large; however, because the entire anal fin lies tiforms, in the goodeids (Uyeno and Miller, farther away from the vertebral column, the 1972) and in New World cyprinodontines dorsal extent of the inclinators is the base of (Uyeno and Miller, 1971; Miller and Walters, the hypaxial musculature. 1972; Levin and Foster, 1972). In both, males 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 131 have fewer chromosomes than females (for goodeids, male 2n = 41, female 2n = 42; for cyprinodontines, male 2n = 47, female 2n = 48), a condition believed caused by centric fusion ofan acrocentric autosome with a similar sex chromosome (Uyeno and Miller, 1972). Heterochromy is apparently present in 0. laucaensis; however, the mechanism for the production ofan increase in chromosome number is unknown (Arratia, 1982). SEXUAL DIMORPHISM AND DICHROMATISM: Killifishes are characterized by being highly sexually dimorphic and dichromatic. Al- though certain aspects of the different patterns are difficult to analyze, Parenti (1981) concluded that females larger than males is the plesiomorphic condition for killifishes. This is true for Orestias in which adult females can dwarf adult males (e.g., 0. parinacotensis, fig. 52). Also, in most species of Orestias where sample sizes are large enough for generalizations, males are far less numerous than females. One species in the gilsoni complex, 0. tchernavini, was described from a holotype and 36 paratypes, all ofwhich are female (Lauzanne, 1981); no males ofthis species have ever been reported. Collections oflarge females in high proportions are characteristic of other cyprinodontiform groups, such as viviparous poeciliids where all female gynogenetic species are known. No experimental work has been published reporting a mode of reproduction in all-female Orestias species. Collections are far too sparse and in the case of older material, lacking in precise FIG. 18. A. Orestias tomcooni (AMNH 52189); locality data, to assess the reproductive bi- B. Orestias elegans (CAS 46179). Diagrammatic ology of Orestias species with high propor- representation of internal caudal fin supports and tions of females. vertebral column. Basal portions ofcaudal fin rays Sexual dichromatism among Orestias blackened. species of the agassii complex follows an ontogenetic pattern comparable to that found in other killifishes; that is, juvenile pigmen- mens that are uniformly dark brown or green- tation patterns persist with little modification ish brown on the dorsal and lateral surfaces, in subadult females, whereas the pattern with the ventrum uniformly pale yellow and changes in adult males and females. the fins clear. The variety was described from Pellegrin (1904b) described four varieties specimens 60 to 110 mm in total length from of 0. agassii, all based on differences in color Lago Titicaca. pattern and size. The four varieties have all Orestias agassii var. typica, so named be- been determined as different life stages in 0. cause Pellegrin believed that it most closely agassii. Each of the four life stages has been resembled the types of 0. agassii, was used identified in one collection from Lago Titi- for specimens 70 to 110 mm in total length caca (fig. 19). Orestias agassii var. inornata that are dark brown or greenish brown on the was the name applied by Pellegrin to speci- dorsal surface, but with a faint mottled pat- 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

:1, zle, -.f.-10' ,-oIt 11.4iY"I"'-ti kh.- 11444 lk

FIG. 19. Orestias agassii (AMNH 52148), from Bahia de Puno, Lago Titicaca. Ontogenetic changes in pigmentation pattern represented by, from top to bottom: adult male, 71.2 mm SL, adult female, 69.9 mm SL, adult female, 57.7 mm SL, subadult male, 41.3 mm SL. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 133 tern elsewhere, and an indistinct dark lateral dividuals of the species listed above; how- body band (fig. 19). Fins are slightly mottled ever, no single lot of any of the species con- rather than clear as in the inornata variety. tained specimens that all exhibited the The third variety, 0. agassii var. senechali, pattern. Whether this is due to individual was distinguished from the typica variety by variation or simply vagaries in preservation having a darker and therefore more distinct and permanence ofthe pigments is unknown. lateral body band and several indistinct dark JAW AND JAW SUSPENSORIUM: Orestias blotches ventral to the band. It was described species have relatively small mouths and up- from specimens 50 to 70 mm in total length. turned lowerjaws. A defining character ofthe The fourth variety, 0. agassii var. crequji genus (fig. 2, character 27) is the absence of (fig. 19) was described from specimens 55 to a ventral medial process in the anguloarticu- 70 mm in total length with mottled dorsal lar bone of the lower jaw. Even with this and lateral surfaces, including an interrupted reduction, the large medial process on the lateral body band. Fins are mottled, rather dentary gives Orestias a very robust lower than clear. jaw. Species ofthe agassii complex pass through Two distantly related species of Orestias ontogenetic changes in color pattern similar (ispi, fig. 34 and jussiei, fig. 50) have lower to that described above for 0. agassii; that jaws that are perpendicular to the body axis. is, juveniles have a relatively mottled, irreg- Other species (e.g., 0. tomcooni fig. 71, and ular pigmentation pattern that is retained with 0. pentlandii, fig. 33) have lower jaws that some modification, in the form ofdarkening, approximate the vertical, but in no two species in subadult females. Adult males are nearly other than ispi and jussiei is the lower jaw so uniformly dark brown or greenish brown with nearly perpendicular to the main body axis. pale yellow ventral surfaces. Very large adult However, a vertical jaw is attained in these females of the complex tend to be more uni- two species in different ways. Orestiasjussiei formly darkened and therefore resemble (fig. 20A) has characteristics of the jaw and males. jaw suspensorium that are considered prim- Species of the cuvieri, gilsoni, and mulleri itive for the genus; 0. ispi (fig. 20B) on the complexes exhibit little or no ontogenetic other hand, has characteristics of the system change in color pattern; adults retain the pat- that uniquely define a group of species in the tern of juveniles. In the gilsoni and mulleri cuvieri complex (ispi, pentlandii, and cuvierl) complexes, adults generally retain the typical as monophyletic. The relatively primitive jaw mottled patterns of juveniles; in the cuvieri andjaw suspensorium characters of 0. jussiei complex, both young and adults are uniform- are described first and then contrasted with ly colored with little mottling, and the dorsal the states of these characters in 0. ispi. surface slightly darker than the ventral. In 0. jussiei (fig. 20A) the premaxilla has A variety of color patterns characterize the S shape characteristic of all cyprinodon- species or groups of species; these are dealt toid fishes (Parenti, 1981, fig. 3b). The as- with in detail in the following sections. One cending process of the premaxilla is narrow species group of the agassii complex is de- and comes to a slender point posteriorly. The fined by a particular pattern of fin pigmen- rostral cartilage is absent in all Orestias. The tation. In these species (frontosus, empy- dentary is a robust bone that carries a sensory raeus, tschudii, richersoni, ctenolepis, and canal. There is an irregular row of unicuspid polonorum) there is a prominent band along teeth on both the premaxilla and the dentary. the base of the pectoral fin rays (fig. 46) ac- The maxilla has a large dorsal process which companied by a faint to dark band at the base has an irregularly grooved surface. of the dorsal and the anal fin rays. These In 0. ispi (fig. 20B), the premaxilla is much pigments are not stable, and are barely dis- wider in a lateral view than injussiei, and the cernible in specimens that have been pre- S shape is much less pronounced. The as- served for a long period of time. In recently cending process of the premaxilla is reduced collected material, the pigments are much slightly and blunt at the posterior tip. The stronger. The pattern is seen best in large in- dentary is a relatively large bone, except at 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

A B

FIG. 20. A. Orestiasjussiei (CAS 46175); B. Orestias ispi (AMNH 52111). Diagrammatic representation of left lateral view ofjaws and jaw suspensoria. the medial extent where it is constricted. siomorphic for all Orestias and present in the There are no discernible troughs for sensory agassii complex, are described for 0. agassii canals in the dentary. The maxilla is a much (fig. 21A) and will serve as a basis of com- more slender bone with a small dorsal pro- parison for more derived skull types within cess that has an unsculptured dorsal surface. the genus. The shapes and relative sizes of the pre- The mesethmoid is present as two large maxilla, dentary, and maxilla are essentially ossified discs that meet anteriorly to form a the same in ispi, pentlandii, cuvieri. The most wedge located at the anterior extent of the striking difference is that the first two species anterior ramus of the parasphenoid (Rosen, have few or no teeth in the jaws (fig. 20B), 1964). The medial extensions of the lateral whereas, quite in contrast, cuvieri (fig. 32) has ethmoids meet or pass dorsal to this ramus. a large mouth with one row oflarge, recurved, The nasals are large, flat bones that lie dor- unicuspid teeth and an irregular row ofsmall- sal to the ethmoid region. They are expanded er, unicuspid teeth interior to the large row. in Orestias to nearly meet in the midline. The As discussed previously in the section on frontals are relatively flat bones that are hyobranchial apparatus, ispi and pentlandii curved ventrolaterally to form the dorsal rim are planktivores, whereas cuvieri is an om- of the orbit. nivore. Gill arch structure varies little among The dermosphenotic is absent. (It is pres- these three species, however. ent as a small ossified element in some spec- There is little difference between the jaw imens of ascotanensis and mundus of the suspensoria ofjussiei and ispi. In both, as in agassii complex.) The dermosphenotic is all Orestias, the autopalatine has a blunt head either absent or present as a minute ossified that is not capped by cartilage. element in Kosswigichthys, contrary to Pa- SKULL: The vomer is absent in all Orestias renti (1981) in which it was stated that the species (fig. 2, character 23); this is the sole dermosphenotic is absent in that genus. synapomorphic skull character of the genus. The first vertebra articulates with the skull Other major features ofthe skull that are ple- via a well-developed basioccipital condyle, 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 135

A B FIG. 21. A. Orestias agassii (BMNH 1862.11.15:3,4); B. Orestias mulleri (AMNH 52118). Diagram- matic representation of ventral view of skull. and the two exoccipital condyles which have series. The data accumulated for these char- weakly formed articulation facets.~~~BOC-acters are summarized in tables 3 through 7, These characteristics of the skull are mod- respectively. ified (i.e., found in a relatively more derived The modal number ofvertebrae is 32, with state) in the other species complexes. In the an observed range of 30 to 38 (table 3). In mulleri, gilsoni, and cuvieri complexes, the three species, the mode is increased: in cuvieri medial extensions of the lateral ethmoids do and tschudii to 34, and in pentlandii to 37. not meet, therefore do not pass dorsal to the These increases are most parsimoniously as- anterior ramus ofthe parasphenoid (fig. 21 B). sessed as autapomorphies ofeach ofthe three The lateral ethmoids are both relatively species because they represent stages of sep- smaller and their medial margins are farther arate transition series from a relatively low apart. number of vertebrae in the outgroup Koss- The orbits project dorsally above the pri- wigichthys which has 28. Two species, taquiri mary dorsal profile ofthe head in the mulleri and minimus, have a mode of 30 vertebrae. and gilsoni complexes (see especially figs. 36, This state may be a stage in a transition series 63, and 69). The orbits are expanded to dif- from a low to a high number of vertebrae. ferent degrees among the 15 species of the Therefore, although useful in distinguishing two complexes, but in all the frontal bones taquiri and minimus from other Orestias are convex along the dorsal rim of the orbit species, it is not considered a synapomorphy to accommodate the relatively large eyes. for the two species. MERISTIC CHARACTERS: Five meristic char- Dorsal fin ray number ranges from 10 to acters were found to be useful in discrimi- 18, with a mode of 14 (table 4). In the sister nating among species or groups ofspecies: (1) species crawfordi and incae there is an in- vertebrae; (2) dorsal fin rays; (3) anal fin rays; crease in the modal number ofdorsal fin rays (4) pectoral fin rays; and (5) scales in a lateral to 16 and 17, respectively. Kosswigichthys 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

TABLE 3 Distribution of the Number of Vertebrae in Orestias Number of Vertebrae Species 30 3 1 32 33 34 35 36 37 38 cuvieri - - - 2 4 4 pentlandi 1 5 1 ispi - 5 19 9 _ forgeti - 1 4 13 5 - mulleri - 4 3 - - - gracl/is _ _ _ 1 - - crawfordi - 10 2 - - - tutini 1 2 - - - - incae 10 - - - luteus - 2 4 1 - - rotundipinnis - 1 2 - - farfani _ _ 1 - - albus - 2 2 1 1 - olivaceus - 1 4 - - - silustani - 1 2 2 - - agassii - 5 5 7 1 3 empyraeus _ 5 1 - - - frontosus - 1 2 - 1 - polonorum - 1 1 - - - elegans - - 3 5 - - jussiei - 1 5 2 - - puni - 1 1 - - - parinacotensisa laucaensisa tschudii - - - 1 3 2 gymnotus - 1 1 1 1 - hardini - - 4 3 - - ctenolepis - - 1 _ _ _ ascotanensis - 2 4 - - - richersoni - - 1 _ _ _ multiporis _ _ 1 - - mundus - - 1 _ _ ututo _ _ 5 _ _ gilsoni - 1 2 - - - taquiri 4 1 - - - - mooni - - 3 3 1 - uruni - 2 1 1 - - minimus 2 - _ _ _ minutus - 1 - _ tchernavini 2 13 10 -- - imarpe - 1 - _ _ tomcooni - 2 - _ _ robustus 1 - _ _

a parinacotensis and laucaensis have not been examined, and no vertebral counts recorded.

asquamatus (BMNH 1948.3.15:40-43) has number of dorsal fin rays to 16 in richersoni 12 dorsal fin rays. Therefore, based on out- is considered to be an autapomorphy of that group comparison, the increase in the former species because on the basis ofall other char- two species is considered to be one of their acters, it is assessed as a distant relative of synapomorphies. The increase in the modal crawfordi and incae. 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 137

TABLE 4 Distribution of the Number of Dorsal Fin Rays in Orestias Number of Dorsal Fin Rays

Species 10 1 1 12 13 14 15 16 17 18

cuvieri - 1 1 1 6 3 pentlandii - 1 1 3 3 3 ispi - 3 22 13 3 - forgeti 2 16 15 6 __ mulleri - 2 7 4 _ _ gracilis 1 2 1 - crawfordi 1 8 2 1 tutini 2 2 incae - 2 2 1 4 1 luteus 1 4 2 1 _ _ rotundipinnis - 3 1 farfani 1 _ _ 1 albus 1 3 5 olivaceus - 1 4 silustani 1 5 2 1 _ agassii - 2 7 7 11 7 2 _ empyraeus - 3 1 4 1 2 frontosus - 2 3 2 1 - polonorum 1 4 - elegans 1 4 6 jussiei 1 7 2 puni 1 2 2 parinacotensis laucaensis - X ------xa tschudii - 1 4 gymnotus - 4 2 1 4 - hardini 1 5 3 ctenolepis _ _ 1 2 5 1 ascotanensis 1 2 3 1 richersoni - 1 1 3 _ multiporis 2 5 3 mundus ~~~~2 5 4 - ututo 3 2 1 gilsoni 2 3 4 1

taquiri - 2 4 1 1 - mooni _ 1 1 1 1

uruni _ 2 2 3 - minimus 2 _ minutus 1 - 1

tchernavini - 4 15 13 5 - imarpe 1 4 1 1 tomcooni _ 1 5 robustus - 1 2 3 3 1

a These data are taken from Arratia (1982) in which dorsal fin ray numbers were presented as a range, rather than frequency distribution.

A reduction in the modal number ofdorsal Anal fin rays number from 10 to 19 with fin rays to 10 in ututo and 11 in hardini is a mode of 15 (table 5). Because a specimen considered to be a synapomorphy of these of Kosswigichthys asquamatus (BMNH two sister species. 1948.3.15:40-43) has 12 anal fin rays, an in- 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

TABLE 5 Distribution of the Number of Anal Fin Rays in Orestias Number of Anal Fin Rays Species 10 11 12 13 14 15 16 17 18 19 cuvieri __ - - 1 2 3 2 2 1 pentlandii - - - - - 2 4 3 1 _ ispl _ _ _ _ - - 15 20 4 2 forgeti _ - -- 2 5 19 1 1 1 1 mulleri - - 3 6 2 1 - gracilis _ _ _ _ 1 3 1 crawfordi _ _ _ 1 7 1 2 _ _ tutini - - - 1 2 1 - incae - - -- 2 1 2 luteus - - - - 2 5 2 rotundipinnis -- - 1 2 3 - farfani - - - - - 1 1 albus - - - - 3 3 3 olivaceus __ _ 1 1 3 silustani - - - - 1 4 4 agassii - 2 5 7 12 6 5 empyraeus - - -- 3 5 2 frontosus - - -- - 6 1 2 _ _ polonorum - - -- 3 1 1 elegans - - -- 1 5 4 jussiei - - - 2 6 2 - puni - - - - 1 1 2 parinacotensis X----X- xa laucaensis - - X------xa tschudii - - - -- 2 5 gymnotus - 2 1 4 2 1 - hardini 1 2 5 - - - - ctenolepis - - - - 5 4 - ascotanensis - - 1 4 4 - -

richersoni _ _ _ _ - 1 3 1 _ multiporis - - - - 2 6 2 mundus -- - 3 6 2 - ututo 2 3 1 - - - - gilsoni - - - 2 2 2 - taquiri - - 1 2 2 1 - moon' _ _ _ 1 2 1 uruni _ _ 1 6 - -

minimus _ _ _ _ 1 1 _

minutus 1 _ tchernavini - - - 6 12 16 3

imarpe - - 2 2 1 1 - tomcooni - - - 1 1 4 - robustus - - - 3 2 2 -

a These data are taken from Arratia (1982) in which anal fin ray numbers were presented as a range, rather than a frequency distribution.

crease in the modal number of anal fin rays modal number of 11 anal fin rays occurs in might be considered a synapomorphy of Or- ututo; the range is from 10 to 12. The de- estias species. However, 12 anal fin rays is crease is considered an autapomorphy of the not an uncommon number in agassii, one of species. Similarly, there is a distinct increase the more variable sDecies. Also. a rather low in the mode to 16 or 17 in the cuvieri com- 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 139

TABLE 6 Distribution of the Number of Pectoral Fin Rays in Orestias Number of Pectoral Fin Rays Species 14 15 16 17 18 19 20 21 22

cuvieri - - 2 2 2 - - - -

pentlandii - - - 1 6 4 1 - -

ispi - 1 7 23 10 - - - - forgeti - - - - 7 18 14 - -

mulleri - - - 5 3 5 - - - gracilis 2 6 ------

crawfordi - - - 1 1 6 1 - -

tutini _ 1 1 1 _ _ _

incae - - 1 2 4 2 1 - - luteus - - - - - 2 4 3 1 rotundipinnis - - - - - 2 3 - - _ farfani - 2 - -

albus - - - - 2 1 2 2 -

olivaceus ------4 1

silustani - - - - 1 1 7 1 -

agassii 1 3 11 9 8 3 1 - - empyraeus - - 4 4 1 - - - - frontosus - - 1 4 1 1 _ _ polonorum - - 3 2 - - - - _ elegans - 3 5 2 1 1 - - - jussiei - 1 4 4 1 - - - - puni _ _ 3 1 1 parinacotensis X ------Xa

Xa laucaensis - X - --

tschudii - - - 1 2 4 3 - - gymnotus 1 3 4 2 1 - - - -

hardini - 4 2 ------

ctenolepis - 1 2 5 1 - - - -

ascotanensis - - - 3 4 1 - - - richersoni - - 1 - 3 1 - - -

multiporis - 1 5 4 3 - - - -

mundus - 1 4 4 3 - - - -

ututo - - - 3 2 - - - - gilsoni - - - 2 2 1 - - -

taquiri - - 1 2 2 - - - -

mooni 1 3 5 1 - - - - - uruni - 1 2 1 - _ _ _

minimus _ _

minutus _ _ _

tchernavini 1 5 6 23 1 1 - - - imarpe - - 1 4 1 _ tomcooni - 3 2 1 - - _ _ _ robustus - - 1 1 4 1 - - -

a These data are taken from Arratia (1982) in which pec toral fin ray numbers were presented as a range, rather than frequency distribution.

plex, considered to be an additional synapo- the modal number to 20 or 21 is a synapomorphic character of the included four morphy of a group of agassii complex species. species. Similarly, an increase in the modal number Pectoral fin rays number from 14 to 22, to 19 is considered an independently ac- with a mode of 17 (table 6). An increase in quired autapomorphy of 0. forgeti of the cu- O II -

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O vieri complex. Pectoral fin rays number 13 or 14 in Kosswigichthys, indicating as for anal fin ray number, that an increase in the modal ON00 number may be a synapomorphy of Orestias species. However, surveys of pectoral fin ray number as well as anal fin ray number among all Anatolian cyprinodontids would be nec- Cl 0u essary to judge at what level the increase in fin ray number should be used as a synapo- *morphy. In the meantime, the low number ofpectoral fin rays in gracilis (range 14 to 15, mode of 15) may be another character to distinguish this species from the rest ofthe mulleri complex (range 15 to 20, mode of 19). The number of scales in a lateral series from the posterior margin of the supracleithrum to the hypural joint varies widely CZ r. among Orestias species as well as among in- 0 dividuals (table 7). The range, when scales C.) CA are present, is from 29 to 60. Juveniles and C._ CZ adults of some species (e.g., minimus) are totally unscaled. The modal number of scales in a lateral series is 32, yet many species have C-) N CZ a wide range (e.g., forgeti, 29 to 39, with a n CO mode of 33), and, tchernavini, 30 to 41, with CZI (Z) C)C.. a mode of 34). Nj (A A synapomorphic increase in lateral scale number occurs in three species of the cuvieri Hl complex (cuvieri, ispi, and pentlandii) in which there is a modal number of41, 48, and IJ 57, respectively. The extremely high number tN in ispi and pentlandii is an additional syn- en CZ apomorphic character of these sister species. cl Other differences in modal number ofscales II - in a lateral series occur between pairs or larger u00C) I QCZ groups of species and will be used in the synapomorphy scheme and key. For example, all members of the mulleri complex have a I. - modal number of 31 or 32, except for mulleri Cl which has a modal number of 36. Also, tom- ON00 cooni of the gilsoni complex has a modal number of37. The extremely high scale num- _ ber in the cuvieri complex is assessed as synapomorphic; however, to judge whether the O -1 1I E high modal numbers of mulleri and tomcooni cl 0 are primitive or derived is confounded by the fact that in Kosswigichthys there are either no CZ scales (as in asquamatus) or, for example, 37 scales (as in a representative of K. splendens, BMNH 1948.3.15:37-39). J) .~ ~ - fi< H A general statement that can be made about meristic variation within Orestias is that it is useful in distinguishing among species or 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

TABLE 8 Frequency Distribution of Length of the Head in Orestias (expressed in thousandths of Standard Length) Class Intervals of Head Length Species 150-174 175-199 200-224 225-249 250-274 275-299 300-324 325-349 350-374 375-399 cuvieri ------2 4 1 1 pentlandii - - 1 2 2 - - - - - ispi - - 5 25 23 3 - - - - forgeti _ 20 5 - - - - - mulleri - - - - 1 4 5 - - - gracilis - - - - 3 2 - - _ _ crawfordi - - - - - 3 5 - - - _ _ _ _ tutini _ - 4 - - - incae - - - - - 3 4 - luteus - - - - - 1 2 3 2 2 rotundipinnis ------3 2 - - farfani ------1 1 - - albus ------3 2 olivaceus - - - - 1 - - 4 - - silustani - - - - 2 1 3 1 - - agassii - - - 6 10 7 2 1 - - empyraeus - - - 1 1 4 2 1 - - frontosus - - - 1 3 - - - - - polonorum - - - 3 2 - - - - - elegans - - - - - 5 3 - - - jussiei _ _ _ - 2 6 - - - puni ------2 2 - - Xa parinacotensis x ....------laucaensis ,,,,,,,,,xa- tschudii - - - - 4 2 - - - - gymnotus - - 1 3 5 1 - - - - hardini - - - - 2 2 3 - - - ctenolepis - - - 3 6 - - - - - ascotanensis - - - - - 1 7 4 - - richersoni _ _ _ _ 5 _ _ _ _ _ multiporis - - - 1 7 2 - - - - mundus - - - - 1 4 5 - - - ututo - - - 2 2 2 - - - - gilsoni - - - - 1 4 3 - - - taquiri - - - - 1 1 3 - - - mooni - - - - 5 2 2 - - - uruni - - - - 1 2 2 - - - _ minimus _ _ - 1 - - - minutus - - - - 1 1 - - - - tchernavini - - - 7 27 3 - - - - imarpe - - - - - 3 1 1 - - tomcooni - - - 5 1 - - - - - robustus - - - - 2 - 4 - - - a These data are taken from Arratia (1982) in which head length was presented as a range, rather than a frequency distribution. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 143

TABLE 9 Frequency Distribution of Width of the Head in Orestias (expressed in thousandths of Standard Length) Class Intervals of Head Width 110- 130- 160- 190- 210- 230- 260- 290- 320- 350- 380- Species 129 159 189 209 229 259 289 319 349 379 409 cuvieri 2 1 1 2 2 - pentlandii 1 2 2 - - - ispi 2 3 2 -- - forgeti 1 3 3 -- - mulleri - - 5 3 - - gracilis 1 3 - - - - crawfordi - - - - 7 2 tutini - - 2 1 1 - incae - - - 1 4 - luteus 2 1 4 4 1 rotundipinnis - 1 1 - farfani 1 - 1 - - _ - - 2 albus - - 3 - _ _ olivaceus - - 1 - - 2 silustani - - 2 5 1 - agassii - 9 15 4 - - empyraeus - 2 3 3 - - frontosus - - 4 2 1 - polonorum - 2 3 -- - elegans - 5 2 - - - jussiei - - - 3 3 4 puni parinacotensis - x ..------xa - - laucaensis tschudii - - 4 1 1 - gymnotus 3 4 2 - - - hardini - - 7 - - - ctenolepis - 5 3 1 - - ascotanensis - 1 2 4 3 - richersoni - 3 2 - - - multiporis - 1 9 - _ _ mundus - 1 5 5 - - ututo - - 5 1 - - gilsoni - - - 7 1 - taquiri - - 3 2 - - mooni - 1 6 1 - - uruni - - 3 2 - - minimus - - 1 - _ _ minutus - - 2 - _ _ tchernavini - 4 2 1 - - imarpe - - 1 4 - - tomcooni 2 2 - - - - robustus - - 1 3 2 -

a These data are taken from Arratia (1982) in which head width was presented as a range, rather than a frequency distribution. 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

TABLE 10 Frequency Distribution of Depth of the Body at Posterior Extent of the Head (expressed in thousandths of Standard Length) Class Intervals of Depth of Head Species 140-169 170-199 200-229 230-259 260-289 290-319 320-349 350-379 380-409

cuvieri 2 2 - 2 pentlandii 2 2 ispi 1 3 4 1 - forgeti 1 4 3 - mulleri 1 4 2 2 - _ _ gracilis 3 crawfordi - 2 2 5 - - tutini - 3 1 - - incae 2 4 - - luteus 2 - 3 4 rotundipinnis 1 3 1 - farfani - 1 - 1 _ _ albus 3 4 - - olivaceus 1 - 3 1 _ _ silustani - 4 2 1 - - agassii 6 9 10 2 1 _ _ empyraeus 4 2 frontosus 3 3 polonorum 2 2 elegans 5 2 1 jussiei 1 3 3 2 - - puni _ 1 2 1 parinacotensis x ------laucaensis x ------Xa tschudii - 3 4

4X gymnotus 4 5 - - hardini 5 2 - ctenolepis 2 6 1 ascotanensis 1 3 3 richersoni - 2 3 multiporis 3 7 - 4 mundus 1 4 5 ututo 2 3 1 gilsoni 2 5 2 taquiri 1 3 1 mooni 5 2 - uruni 1 2 2 minimus _ 1 _ minutus 1 1 - tchernavini 5 4 - imarpe - 4 1 tomcooni 2 1 - robustus 1 1 4 a These data are taken from Arratia (1982) in which head depth was presented as a range, rather than a frequency distribution. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 145

TABLE 11 Frequency Distribution of Depth of the Caudal Peduncle (expressed in thousandths of Standard Length) Class Intervals of Depth of Caudal Peduncle Species 050-074 075-099 100-124 125-149 150-174 175-199 200-224 cuvieri - 1 3 2 pentlandii 3 2 ispi - 26 21 2 forgeti 15 31 mulleri 1 6 gracilis 3 1 crawfordi 3 5 2 tutini 3 incae 1 4 luteus 3 3 2 rotundipinnis 3 farfani 1 albus 3 2 olivaceus 15 4 silustani 15 2 agassui 10 2 empyraeus 2 3 3 frontosus 5 polonorum 4 elegans 3 2 jussiei 1 4 puni 5 3 parinacotensis laucaensis x - .------xa tschudii 2 2 gymnotus - 2 3 4 hardini 6 ctenolepis 7 2 ascotanensis 3 5 2 richersoni _ 5 multiporis 4 6 mundus 2 5 ututo 2 2 gilsoni 7 2 taquiri 5 mooni 1 2 uruni 3 2 minimus minutus 1 tchernavini - 2 35 imarpe 4 tomcooni 1 2 1 robustus 2 4

a These data are taken from Arratia (1982) in which depth of the caudal peduncle was given as a range, rather than a frequency distribution. 146 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

TABLE 12 Frequency Distribution of Preorbital Distance (expressed in thousandths of Standard Length) Class Intervals of Preorbital Distance Species 020-039 040-059 060-079 080-099 100-119 120-139 cuvieri - - 2 3 3 pentlandii 2 4 - - ispi - - 3 4 1 forgeti - 4 1 - mulleri - 5 4 - gracl/is - 4 _ _ crawfordi 2 4 3 - tutini - 3 1 - incae - - 4 2 luteus 2 4 4 - rotundipinnis - 2 1 - farfani - 2 - _ albus - - 2 5 olivaceus - 2 3 - silustani 2 4 3 - agassia 1 20 6 - empyraeus - 1 5 4 frontosus - 4 2 - polonorum _ 5 _ _ elegans - 3 2 - jussiei - 4 5 - puni _ _ 3 - parinacotensisr laucaensisa tschudii - - 2 2 - gymnotus - 3 5 1 hardini - 3 4 - ctenolepis ascotanensis - 3 6 - richersoni 1 4 - - multiporis - 8 2 - mundus 1 6 3 - ututo - - 5 1 - gilsoni - 8 1 - taquiri - 2 2 1 mooni - 5 2 - uruni - 4 1 - minimus _ _ 1 - minutus 1 - 1 - tchernavini 3 2 _ _ - imarpe - 4 1 - tomcooni 2 2 - - robustus 1 4 1 - a No data available for these species. species groups, and that obvious synapo- MORPHOMETRIC CHARACTERS: Five addi- morphies can be used to define groups of tional characters that are useful for distin- species; but, determination of polarity of in- guishing among Orestias species and that creasing or decreasing modes is problematic. are best described as morphometrics are: 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 147

70W 619°W

PERU BOLIVIA

Az'angarO I~~~~~~~~ _QB ) de~~~~~~~~~~~~~~~~~~~~~~~~~~U La na rapa~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~r

¶ Huancane

R. Mhoh[o

Umayo l de zd Puno oALAGO \GRANDE

C/) 0~~~~~~~~~~~~~~~~~~~~~

C,,~~~~~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~,'EQ~ A ~~~~~~LG

Desaguadero Tihunaco

/ 0~~~~~~~~~~~~~~ <(#>o 710W 692 FIG. 22. Major divisions of Lago Titicaca, surrounding towns and tributaries. Labeled islands are: A. Soto, B. Chiquipa, C. Amantani, D. Taquile, E. Titicaca, F. Coati, G. Taquiri, H. Paco. 148 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

Peru Bolivia

O multi p or is * ncae * r i c h e r s o n O c raw f ord'i

_ moon1 0 mrinutus,uruns * s'lustani, ol ivaceus

1+ pun

O r ot uncd ip in n is

,-" robustus al minimus ctenolep is farfani

O forget i,tch iernavini, ispi

tut c 'ilsoni, taquiri A ini, O tomcoon i, imarpe, gracilis i * muller i / O agassii

70 6,9 0

FIG. 23. Type localities within the Titicaca Basin. For names of sections of Lago Titicaca, towns, tributaries and islands, see figure 22. Not represented by a symbol because type locality is no more specific than "Lago Titicaca": cuvieri, pentlandii, luteus, albus, tschudii, and frontosus. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 149

FIG. 24. Type localities north of the Titicaca Basin.

(1) length of the head, from tip of the lower as synapomorphies of species groups be- jaw to the posterior extent of the operculum; cause, as will be discussed, all of these char- (2) width ofthe head taken across the middle acters are not correlated. of the opercula; (3) depth of the body taken The head is very large reaching nearly 40 at the posterior margin ofthe operculum; (4) percent of SL in four species: cuvieri of the depth of the caudal peduncle; and (5) preor- cuvier complex and luteus, rotundipinnis and bital distance. The variation of these char- albus of the agassii complex (table 8). This acters, expressed in terms of a sixth variable, character is autapomorphous for cuvieri and SL, is summarized in tables 8 to 12, respec- for albus, and an additional synapomorphic tively. All are straight line measurements re- character of the two sister species, luteus and corded with metric dial calipers. rotundipinnis. Tchemavin (1944a, p. 147) pointed out that The head (table 9) of the three species, lu- with an increase in size (i.e., SL) in Orestias teus, rotundipinnis, and farfani, attains a species, there is an increase in the depth of width of over 34 percent of SL. In some in- the caudal peduncle, length ofthe head, width dividuals of luteus, the head is wider than it ofthe head, length ofthe snout and correlated is long. Two other species of the luteus group changes in a variety of other morphometric (olivaceus and albus) have heads that are wid- characters. This is true as a general statement; er on the average than other Orestias but not however, morphometric characters are used so wide as in the three aforementioned species. 150 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

in the depth of the body within the mulleri complex, with crawfordi, tutini, and incae reaching a depth of over 34 percent of SL. Being deep-bodied is also a synapomorphy of a group within the agassii complex; that is, species of the luteus group and 0. puni and jussiei. Depth of the body (table 10) is correlated in a number of species with depth of the caudal peduncle (table 1 1). On the average, all members of the agassii complex have a relatively deep caudal peduncle. A deep caudal peduncle occurs also in crawfordi and incae, two relatively deep-bodied species of the mulleri complex. By comparison with Kosswigichthys, an elongate, narrow-bodied genus, the increase FIG. 25. Type localities in Chile. in depth of the body and of the caudal peduncle is most parsimoniously assessed as independently derived in the mulleri and in A comparison of tables 8 and 9, shows that the agassii complexes. length ofthe head is not correlated with width The preorbital distance (from the tip ofthe of the head, as cuvieri has a head of average lower jaw to the anterior rim of the orbit) width. reaches nearly 14 percent of SL in cuvieri Orestias luteus and rotundipinnis are ex- (table 12). This derived character can also be tremely deep-bodied (table 10) as are three described as an increase in the length of the other species ofthe luteus group (farfani, oli- lower jaw (fig. 32) which is the longest in the vaceus, and albus). Also, there is an increase genus in cuvieri.

EXPLANATION OF SYNAPOMORPHY DIAGRAMS A single diagram depicting the relation- The following synapomorphies are those ships and distribution of synapomorphic described in detail in the previous section, or characters among the 43 species of Orestias are additional, newly described data, supple- would be a confusing and cumbersome figure. mented by data from Allen, in Eigenmann Therefore, six synapomorphy diagrams are and Allen (1942), Tchemavin (1944a, 1946), presented: (1) a diagram of relationships Lauzanne (198 1), and Arratia (1982). among the four species complexes and the defining characters of each monophyletic RELATIONSHIPS AMONG THE FOUR SPECIES complex (fig. 26); individual diagrams of re- COMPLEXES (fig. 26): The cuvieri, mulleri, and lationship among, and the defining characters gilsoni complexes together form a monophy- ofspecies within, the (2) cuvieri complex (fig. letic group defined by three synapomorphies. 27), (3) mulleri complex (fig. 28), (4) gilsoni (28) Epipleural ribs are strongly bifid in complex (fig. 29), (5) agassii complex, ex- adults, as opposed to being simple in the cluding species of the luteus group (fig. 30), agassii complex. In just one species of these and (6) luteus group (fig. 31). three complexes, forgeti, the ribs are only The numbering of synapomorphic char- weakly bifid. acters continues in sequence from figure 2, (29) Lateral ethmoids are relatively far apart in which the 20 characters used in discussing (fig. 21 B), rather than being close to the para- relationships of cyprinodontids are num- sphenoid (fig. 21 A), and their medial exten- bered 1 through 20, and the seven defining sions do not meet or pass dorsal to the an- characters of Orestias are numbered 21 terior ramus of the parasphenoid. Some through 27. species of the agassii complex (e.g., hardini) 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 151

approach the latter condition; their phylo- x x x x 0. 0 genetic position will be discussed along with 0 E 0 E 0 E E the relationships within that complex. 0 0 0 l (30) There is no marked ontogenetic change Il cc0 z in color pattern, with adults showing little or w w en- -J 0 no sexual dichromatism. (Orestias mundus : -l .) -J 0 of the agassii complex is defined by its uni- U D a form, light brown color with a dark median 38 139 40 43 137 42 line in all specimens examined. However, be- 36 41 cause ofthe inferred relationships of mundus 35 to other agassii complex species, the pattern 34 is considered to be independently derived in mundus.) ~ i3123 The mulleri and gilsoni complexes together share three synapomorphies.

(31) Orbits project dorsally above the pri- p 11pr --30 mary dorsal profile of the head, and the eyes are much larger than in other complexes. To a certain degree, the eyes project more dor- I21-27 sally in the gilsoni complex, although the extent is difficult to quantify. Therefore, this possibly further derived state is not used as FIG. 26. Synapomorphy diagram of cuvieri, an additional defining character ofthe gilsoni mulleri, gilsoni and agassii complexes. Characters complex. are numbered as those in the text. Black squares (32) Longitudinal axis of base of anal fin represent synapomorphies, open squares symple- not parallel to the vertebral column, but rath- siomorphies. er is angled anteriorly away from the body axis (fig. 4B), fin base projecting slightly beyond primary ventral profile, creating a dis- number for the genus is 15 (table 5). The tinct transition between abdomen and caudal outgroup genus Kosswigichthys has 12 anal peduncle. fin rays; therefore, the increase is considered (33) The second pharyngobranchial is re- to be derived within the cuvieri complex. duced to a U-shaped bone, and bears a tooth- mulleri complex plate with few or no teeth (fig. 1 1). (39) Base of the dorsal as well as that of SYNAPOMORPHIES OF FOUR SPECIES COM- the anal fin projects beyond the primary body PLEXES profile (figs. 36 to 40), and usually covered cuvieri complex with small, thin scales. (34) The ventral branchial apparatus is gilsoni complex elongate, including the basihyal which is nar- (40) The procurrent caudal fin rays lie row (fig. 15) rather than being triangular (fig. mostly interior to the body profile and there- 13) or more rectangular (fig. 14), as in the fore closer to the vertebral column (fig. 1 8A), three other complexes. giving the caudal fin a tapered external ap- (35) Fifth ceratobranchials (lower pharyn- pearance. geals) are narrow and very close together or fused along the midline in adults (fig. 15). agassii complex (36) Dorsal gill arches are reduced relative (41) Lateral scales, from the operculum to to the size of the ventral gill arches. a point at about the posterior extent of the (37) Interarcual cartilage a long, thick rod, pectoral fin, are enlarged with concentric striae equal to or longer than the first epibranchial absent or present to a limited degree. This bone (fig. 10). so-called lateral shield is present in a modi- (38) There is an increase in the modal num- fied form in gymnotus, elegans, ascotanensis, ber of anal fin rays to 16 or 17, with an ob- parinacotensis, and laucaensis as well as utu- served range of from 14 to 19. The modal to, hardini, and mundus. These modifica- 152 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

a a z cc cn 0 _cc -c w w C.)-J w 0I- z z .4 c- -J w a: 4U U 0 z I- 0. l Ui. (. a 155 57 59 161 77 175 73 72 p69 '54 56 58 160 71 F76 '74 '53 170 152 151 , 168 n IofAT i L -A- ::X6

3 ______41___ 1499 48 64 III-C 47 I 1I b46 AlR u ..5"45

34-38 39

FIG. 27. Synapomorphy diagram of species of FIG. 28. Synapomorphy diagram of species of cuvieri complex. Characters are numbered as those the mulleri complex. Characters are numbered as in the text. Black squares represent synapomor- those in the text. Black squares represent synapo- phies, open squares symplesiomorphies. morphies, open squares symplesiomorphies.

tions are discussed in the explanation of the scales ofthe agassii complex is considered as agassii complex synapomorphy diagram (fig. another of its synapomorphic features. 30). Relationships among species of the cuvieri (42) Caudal peduncle is relatively deep in complex (fig. 27): The four species of the cu- adults, reaching over 22 percent ofSL. (There vieri complex (cuvieri, pentlandii, ispi, and is an increase in the depth of the caudal pe- forgeti) are defined as a monophyletic group duncle in two sister species of the mulleri by the five synapomorhies (characters 34 complex, incae and crawfordi, which is as- through 38) discussed above for figure 26. sessed as an independently derived character Orestias cuvieri, pentlandii, and ispi form because ofthose species' close relationship to a monophyletic group defined by four syn- the other species of the mulleri and gilsoni apomorphies. complexes.) (44) There is an increased modal number (43) Scales on dorsal portion of head, par- of scales in a lateral series to 41 in cuvieri, ticularly those ofthe median ridge, enlarged, 48 in ispi, and 57 in pentlandii, with an ob- and usually with no or few discernible con- served range of from 40 to 60 (table 7). centric striae. A similar condition is found in (45) Caudal fin is lunate, rather than round- some large adults of cuvieri and pentlandii, ed or truncate, and the caudal peduncle is although rarely in the latter species. Also, in constricted at the base of the caudal fin (figs. 32 to these two large species ofthe cuvieri complex 34). (46) Premaxilla with a blunt ascending pro- the dorsal scales are never as large relative to cess, and much wider in a lateral view and those lateral to the median dorsal ridge as in the S shape less pronounced (fig. 20B) than the agassii complex. In ispi and forgeti and in other Orestias species (fig. 20A). species of the mulleri and gilsoni complexes, (47) Dentary is a robust bone that is con- a median dorsal ridge is distinguishable, yet stricted near its medial end (fig. 20B). these scales are not thickened, nor smooth or Orestias pentlandii and ispi are defined as granulated. Therefore, the large, smooth sister species by three synapomorphies. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 153

(48) There are few or no teeth in the outer fordi, and incae) are defined as a monophy- jaw in adults (fig. 20B); juveniles almost al- letic group by character 39, base ofthe dorsal ways have a few simple teeth. as well as that of the anal fin extends beyond (49) There are six branchiostegal rays, with the primary dorsal and ventral profiles, re- just one ray anterior to or in the notch on the spectively. anterior ceratohyal (fig. 6A). Orestias mulleri, tutini, crawfordi, and in- (50) There is a further increase in the modal cae form a monophyletic group defined by number of scales in a lateral series to 48 in two synapomorphies. ispi and 57 in pentlandii, with an observed (62) There is an increase in the depth of range of from 41 to 60 (table 7). the body to 23 percent or more of SL (table 10). Autapomorphies of the four species: (63) Body is fully scaled with thin, small 0. cuvieri scales, except in tutini which, although pos- (51) Mouth is extremely large, and there sessing small, thin scales, is not fully scaled. are large, recurved unicuspid outer jaw teeth It is impossible to determine whether the state (fig. 32). in tutini represents the primitive condition, (52) There is an increase in the modal num- or if tutini has secondarily lost scales. ber ofvertebrae to 34 with an observed range Orestias tutini, crawfordi, and incae form of from 33 to 35 (table 3). a monophyletic group defined by two syn- (53) The head is relatively long, its length apomorphies. reaching nearly 40 percent of SL (table 8). (64) There is a further increase in the depth (54) Preorbital distance is greater than in of the body to over 26 percent of SL (table all other Orestias, reaching nearly 14 percent 10). of SL (table 12). (65) The body is laterally compressed, rath- (55) This is the largest species in the genus, er than being relatively fusiform. attaining an observed maximum SL of 220 Orestias crawfordi and incae are defined as mm. sister species by three synapomorphies. (66) There is an increase in the modal num- 0. pentlandii ber of dorsal fin rays to 16 in crawfordi, with (56) There is an increased modal number an observed range of from 15 to 18, and to of vertebrae to 37, with an observed range of 17 in incae, with an observed range of from from 36 to 38 (table 3). 14 to 18 (table 4). (57) There is a further increase in the num- (67) The depth of the caudal peduncle is ber of scales in a lateral series, with a modal increased to 15 percent or more of SL (table number of 57 and an observed range of from 11). 51 to 60 (table 7). (68) Pharyngeal teeth are molariform (fig. 0. ispi 11), although they are consistently broader (58) The main axis of the lower jaw is at a and more blunt in the lower pharyngeals of 90-degree angle to the body axis. crawfordi (fig. 16). (59) The body is fully scaled, with very small, thin, irregularly distributed scales. Autapomorphies of the five species: 0. forgeti 0. gracilis (60) There is an increase in the modal num- (69) The modal number ofpectoral fin rays ber of pectoral fin rays to 19, with an ob- is reduced to 15, with an observed range of served range of from 18 to 20 (table 6). from 14 to 15, as opposed to a modal number (6 1) The body is fully scaled with relatively of 17 for the genus (table 6). small, thin scales, but not as small as those of i. 0. mulleri isp (70) The modal number of scales in a lat- Relationships among species ofthe mulleri eral series is increased to 36, with an observed complex (fig. 28): The five species ofthe mul- range of from 33 to 38 (table 7). leri complex (gracilis, mulleri, tutini, craw- (7 1) Maximum SL obtained (92 mm) is the 154 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

2 CD, z 4 CO Cl,n 0 z n z 0 0. 0 0 z w cc co 0 m a z 0 z 0co -1 4 I- 5- i 86 88 190 92 I94 196 98 I- 1101 103 87 189 1 91 93 95 97 1100 102

80I 82 179 81 Lzz 83

Xi 78

FIG. 29. Synapomorphy diagram of species of gilsoni complex. Characters are numbered as those in the text. Black squares represent synapomorphies.

greatest within the mulleri and gilsoni com- (77) Males with four to five irregular me- plexes. dium to dark brown bars on a lighter brown (72) The modal number of anal fin rays is background; females with fainter bars (all reduced to 13, with an observed range offrom specimens faded in preservation). 12 to 15 (table 5). Relationships among species ofthe gilsoni complex (fig. 29): the 10 species ofthe gilsoni 0. tutini complex are defined as monophyletic by the (73) Dorsal surface of the head, and oper- unique structure of the caudal fin described culum ofmales with fleshy, tubercle-like pro- as character 40 (fig. 26). jections, covering the ctenii of underlying Before presenting the synapomorphies scales. Basal portion ofdorsal, anal, and pec- which describe the relationships among toral fins, and either side of median dorsal species ofthe gilsoni complex, a comment on ridge unscaled; all scales of males apparently several of the species is required. Two, mi- ctenoid (all specimens partially descaled). nutus (fig. 68) and minimus (fig. 67) were 0. crawfordi described by Tchemavin (1944a) from two (74) Fourth epibranchial bone with a huge specimens each. The trivial names of both lateral flange (figs. 11 and 12), and associated species are in reference to the small size of large adductor IV that fills the cupped surface mature adults: minimus was described from of the bone. two gravid females, 23.0 and-28.0 mm in SL; (75) Pharyngeal teeth (fig. 16) are consis- minutus was described from two gravid fe- tently broader and therefore may be de- males, 24.0 and 28.0 mm. Tchemavin (1 944a) scribed as more molariform than in incae. assumed that adult males, if found, would be markedly smaller, following the trend exhib- 0. incae ited by other Orestias species. (76) The most laterally compressed fishes One problem with determining the unique- in the genus Orestias, with the ventral surface ness of maturity at a small size (i.e., SL) is of largest adults nearly keeled. that collections of other, apparently closely 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 155 related species, also from the Percy Sladen Autapomorphies of the 10 species: Titicaca Expedition, were eviscerated for parasite study and their gender or state of Orestias tchernavini sexual maturity cannot be determined. It is (86) The head is relatively laterally com- generally assumed that fishes decline in rate pressed, with a width as low as 13 percent of of increase of SL at maturity. However, this SL (table 9). phenomenon should be further investigated 0. imarpe with laboratory populations to determine its (87) Lips thick and fleshy (fig. 70). generality, especially in killifishes. Therefore, (88) There is a reduction to 1 1, in the mod- it is only tentatively that I accept the unique- al number ofdorsal fin rays, with an observed ness of maturity at a small size in minimus range of from 10 to 14 (table 4). and minutus. Orestias gilsoni, taquiri, minimus, minu- 0. uruni tus, tomcooni, and mooni form a monophy- (89) Basal portion of the pectoral fin is at letic group defined by one synapomorphy. the posterior margin ofthe operculum which (78) Small fishes with an overall body shape obscures the pectoral fin base, rather than characterized by a robust head and relatively being farther posterior, hence separated from narrow caudal peduncle (see tables 10 and the operculum and exposing the pectoral fin I 1) creating an abrupt transition between ab- base. domen and caudal peduncle. (90) Anal pouch and papilla are much re- Orestias gilsoni and taquiri are defined as duced or absent. sister species by two synapomorphies. 0. robustus (70) There is an increase in the modal num- (9 1) Body very robust overall; width ofthe ber of pectoral fin rays to 17 or 18, with an head reaches nearly 23 percent of SL (table observed range of from 16 to 19 (table 6). 9). (80) Pigmentation pattern characterized by (92) There is a gentle rise in the dorsal a silvery background with from six to nine profile from the tip of the snout to the base irregular bars (see figs. 63 and 64). ofthe dorsal fin, and a gentle downward slope Orestias minimus and minutus are defined in the ventral profile from the tip ofthe lower as sister species by two synapomorphies. jaw to the base of the anal fin (fig. 72). (81) Maturity of adults (only females known) is reached at a small size (23.0 mm 0. gilsoni SL in minimus and 24.0 mm SL in minutus). (93) The head is relatively wide and greatly (82) There is a reduction in the number of flattened dorsally, its width reaching nearly scales in a lateral series to 30 or less (minimus 23 percent of SL (table 9). is unscaled; table 7). (94) Caudal peduncle is relatively narrow; Orestias tomcooni and mooni are defined its depth is less than 10 percent ofSL in most as sister species by one synapomorphy. specimens (table 1 1). (83) Main axis of lower jaw is at an abrupt 0. taquiri (nearly 90-degree) angle to the body axis. (95) There is a reduction to 12 in the modal Orestias tchernavini, imarpe, uruni, and number of dorsal fin rays, with an observed robustus form a monophyletic group defined range of from 11 to 14 (table 4). by one synapomorphy. (96) There is a reduction in the modal (84) Pigmentation pattern is characterized number of vertebrae to 30, as opposed to a by a light cream to yellow background cov- mode of 32 for the genus as a whole, and a ered with dark melanophores arranged in a modal number of 31 for the gilsoni complex distinctive marbled pattern (figs. 66, 69, 70, with an observed range offrom 30 to 31 (table and 72). 3). Orestrias imarpe, uruni, and robustus form a monophyletic group defined by one syn- 0. minimus apomorphy. (97) The body is totally unscaled. (85) Robust body, with the depth of the (98) There is a reduction in the modal caudal peduncle reaching nearly 15 percent number of vertebrae to 30 (see character 96 of SL (table 11). above for 0. taquiri). 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

Ul) U)- - z z LU

ozen 4z lx Ur) 4 4c .

FIG. 30. Synapomorphy diagram of species of agassii complex. For relationships within the luteus group, see figure 31. Characters are numbered as those in the text. Black squares represent synapomorphies.

0. minutus agassii complex are defined as a monophy- (99) There is an increase in the number of letic group by the three synapomorphies dis- anal fin rays to 16 or 17, as opposed to a cussed for figure 26 (characters 41 through mode of 15 (table 5). 43). Species of the agassii complex are divided 0. tomcooni into three groups on the basis of squamation (100) The modal number of scales in a lat- patterns described below. Each pattern is as- eral series is increased to 37, with an observed sessed as a uniquely derived condition, and range of from 36 to 38 (table 7). therefore as a defining character for one of (101) Distinctive body shape characterized the three groups. The patterns do not appear by a relatively narrow (table 9) and deep head to be part of a recognizable transition series, (table 10), with an even more abrupt tran- although further data on ontogeny of squa- sition between the anterior part of the body mation patterns might lead to a resolution of and the caudal peduncle (fig. 71) than is di- the trichotomy at the base of the cladogram agnostic of the group comprising gilsoni, ta- of figure 30, by supporting a transition series quiri, minimus, minutus, tomcooni, and hypothesis for these characters. mooni. Orestias ututo, hardini, and mundus form 0. mooni a monophyletic group defined by one syn- (102) A fully scaled species, with very small, apomorphy. thin irregularly distributed scales covering the (104) Lateral scales, those of the so-called ventrum and the basal portion of the pecto- lateral shield, are larger than surrounding ral, dorsal, and anal fins. scales more so than in other agassii complex (103) There is an increase in the modal species. Lateral shield scales overlap to a great number of vertebrae to 32 or 33, with an degree, such that only the posterior margins observed range of from 32 to 34 (table 3). of scales are visible. Relationships among species ofthe agassii Orestias ututo and hardini are defined as complex (figs. 30 and 31); the fishes of the sister species by one synapomorphy. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 157

(105) There is a decrease in the modal (I) z number of dorsal fin rays to 10 in ututo and 2 11 in hardini, with an observed range in both 0: u) of from 10 to 12 (table 4). cn c z LU D z 4 Orestias gymnotus, elegans, ascotanensis, - IL Ca)m U parinacotensis, and laucaensis form a mono- -j -J J 0r U 0 phyletic group defined by one synapomor- 140 1144 K138 1139 4' 143 UP phy. -i c 1142 (106) Body irregularly scaled, and in a par- 141 ticular pattern. Unique characteristics of the pattern concern the configuration of the lat- 127 eral scales: from the posttemporal, scales run l- )126 posteriorly in one or two uneven rows. The 1125 number of scale rows increases posteriorly to 1124 I1 become a series ofapproximately four to five T Th 123 rows at the base of the caudal fin. Dorsal scales may be irregularly and sparsely dis- 1122 tributed. Typically, there are no scales be- H -i '121 neath the pectoral fin (fig. 55). (Specimens of parinacotensis and laucaensis have not been FIG. 31. Synapomorphy diagram of species of examined; a description of these species' ir- luteus group of the agassii complex. Characters regular squamation pattern [Arratia, 1982] are numbered as those in the text. Black squares agrees in many respects with that given above, represent synapomorphies. although is not complete enough to determine if the pattern is identical with that of gymnotus, elegans or ascotanensis. See Sys- tematic Accounts for further remarks.) well as lateral body scales, enlarged and thick- Orestias gymnotus and elegans are defined ened, especially in larger individuals, with as sister species by one synapomorphy. few or no concentric striae, not overlapping (107) Body relatively elongate, with a cor- in larger individuals and usually interdigi- respondingly relatively slender body, the tating along their perimeters. Lateral scales depth of which is as low as 20 percent of SL may have scattered flecks of dark pigment (table 10). (e.g., fig. 48). These head and lateral scales Orestias ascotanensis, parinacotensis, and cannot be removed easily, and give these fish- laucaensis are defined as a monophyletic es an armored appearance. group by one synapomorphy. Orestias ctenolepis, frontosus, tschudii, (108) Extent of squamation sexually di- richersoni, polonorum, and empyraeus form morphic, with females tending to be more a monophyletic group defined by one syn- fully scaled than males; 0. ascotanensis fe- apomorphy. males, for example, have several complete (111) Pigmentation pattern characterized lateral scale rows, whereas males have one by a dark band at the base ofthe dorsal, anal row anteriorly, and several posteriorly. and particularly the pectoral fin rays (see, e.g., Orestias jussiei, puni, agassii, silustani, fig. 46). Pigments are not stable and are barely multiporis, luteus, farfani, rotundipinnis, al- discernible in specimens that have been pre- bus, olivaceus, ctenolepis, frontosus, tschudii, served for a long period of time. richersoni, polonorum, and empyraeus form Orestiasfrontosus, tschudii, and richersoni a monophyletic group defined by two syn- form a monophyletic group defined by one apomorphies. synapomorphy. (109) Anterior ribs become enlarged and (1 12) The modal number ofdorsal fin rays swollen with age (i.e., with increased SL) to is increased to 15 or 16, with an observed create rather wide-bodied individuals (see figs. range of from 14 to 17 (table 4). 7 and 8). Orestias tschudii and richersoni are defined (1 10) Scales on dorsal surface of head, as as sister species by one synapomorphy. 158 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

(113) Body laterally compressed, especially Orestias olivaceus, albus, farfani, rotundi- posterior to the origin of the dorsal fin. pinnis, and luteus (the luteus group; fig. 31) Orestiaspolonorum and empyraeus are de- form a less inclusive monophyletic group defined as sister species by one synapomorphy. fined by one synapomorphy. (114) Median dorsal ridge extremely irreg- (121) Scales from the snout to the base of ular and incomplete, formed by small, smooth the dorsal fin, the preopercular and opercular scales. regions, and dorsal to the pectoral fin are thick, Orestias agassii, jussiei, silustani, puni, oli- and covered with dense granulations (fig. 7). vaceus, albus, rotundipinnis, farfani, and lu- Orestias albus, farfani, rotundipinnis, and teus (the last five species forming the luteus luteus form a less inclusive monophyletic group) form a monophyletic group defined group defined by one synapomorphy. by one synapomorphy. (122) Head at a point just dorsal to the (115) In the largest adults, depth of the operculum is swollen; that is, the scales are body reaches nearly 35 percent of SL (table extremely large and thick, and give the body 10), and is typically more than 29 percent. a rather angular appearance (fig. 8). This (However, agassii is a variable species, and swelling is slightly less pronounced in young large adults are not represented in all collec- individuals, particularly those of albus. tions. See Systematic Accounts for further Orestias farfani, rotundipinnis, and luteus remarks.) form a monophyletic group defined by two Orestias jussiei, silustani, puni, olivaceus, synapomorphies. albus, farfani, rotundipinnis, and luteus form (123) Head is very wide, reaching a width a less inclusive monophyletic group defined of more than 34 percent of SL (table 9). by one synapomorphy. (124) Scales on the operculum are thick (1 16) Width of the head is on the average and granulated, such that the operculum ap- 19 percent or more of SL (table 9). (Orestias pears to be swollen, and arched slightly away puni, jussiei, silustani, and olivaceus do not from the head (fig. 8). have the pronounced shoulders of albus, far- Orestias rotundipinnis and luteus are de- fani, rotundipinnis, and luteus; this character fined as sister species by three synapomor- easily separates the wide-bodied species. See phies. character 122, below.) (125) Head is wider than in any other two Orestias silustani, puni, olivaceus, albus, Orestias species, reaching nearly 38 percent farfani, rotundipinnis, and luteus form a less of SL in rotundipinnis, and 41 percent of SL inclusive monophyletic group defined by one in luteus (table 9). synapomorphy. (126) Pharyngeal teeth are molariform, with (1 17) Pectoral fins are relatively large, the large, rounded cusps. (The state of this char- modal number of fin rays is increased to 20, acter is questionable in farfani.) with an observed range offrom 18 to 22 (table (127) Fifth ceratobranchials (lower pha- 6). ryngeals) are closely apposed, although not Orestias puni, olivaceus, albus, farfani, ro- fused. tundipinnis, and luteus form a less inclusive monophyletic group defined by three syn- Autapomorphies of each of the 24 species: apomorphies. 0. ututo (118) Dorsal profile is arched just posterior (128) There is a decrease in the modal to the head (figs. 7, 41 through 44) creating number of anal fin rays to 11, with an ob- an abrupt transition, in the dorsal profile, be- served range of from 10 to 12 (table 5). tween the head and body. (119) Body is relatively deep, reaching a 0. hardini depth ofjust over 40 percent ofSL (table 10). (129) When a complete series of scales in (120) Head is relatively wide, reaching a a lateral series is present, there is an increase width of over 28 percent ofSL in puni, albus, in the number to a range of from 36 to 38 and olivaceus, and more than 34 percent of (table 7). SL infarfani, rotundipinnis, and luteus (table (130) Squamation incomplete, with dorsal 9). surface of some specimens, and ventral sur- 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 159 face ofall specimens, devoid ofscales. Lateral correspondingly, a body that is not so deep series often incomplete. as in albus, rotundipinnis, and luteus (table 0. mundus 10). (131) Juveniles through adults all uniform- 0. albus ly light brown, with a thin, median dark line (141) Lower jaw very robust, creating a through scales of the lateral series. prominent "chin" (fig. 43). 0. gymnotus (142) Head relatively long, more than 35 (132) Body narrow and elongate, nearly fu- percent of SL in adults (table 8). siform in largest adults (fig. 55) as exemplified (143) Dorsal body profile slopes very gently by a relatively narrow head, width nearly as posteriorly, with no abrupt transition, from low as 10 percent of SL (table 9), as well as the anterior base of the dorsal fin to the base a low depth of the body (table 10). of the caudal fin, the rays of which flare out slightly at its base (fig. 43). 0. elegans (133) There is an increase to 35 or 36 in 0. olivaceus the modal number ofscales in a lateral series, (144) Orbits are oriented dorsolaterally (fig. with an observed range offrom 34 to 38 (table 44), such that in dorsal view, both orbits are 7). nearly completely visible. 0. ascotanensis 0. silustani (134) A robust species, with a relatively (145) Dorsal head and body profile very long head, reaching nearly 35 percent of SL nearly straight from the tip of the snout to (table 8). the anterior base of the dorsal fin (fig. 45). 0. parinacotensis (135) Straight dorsal head profile and head 0. jussiei greatly flattened in both males and females (146) Lower jaw is at a 90-degree angle to the body axis (fig. 50). (fig. 52). (147) Dorsal profile convex, rising very 0. laucaensis sharply from tip ofsnout to middle ofmedian (136) Diploid number of chromosomes is dorsal ridge (fig. 50). increased to 52 in males and 50 or 51 in females, as opposed to a plesiomorphic dip- 0. agassii loid number of 48 in males and females (Ar- (148) Head squamation irregular, usually ratia, 1982). with no head scales anterior to the orbit, and scales absent from either side of the median 0. puni dorsal ridge in young individuals. (See Sys- (137) Caudal peduncle is very deep, reach- tematic Accounts for further remarks on the ing a depth of more than 22 percent of SL definition of this variable species.) (table 11; see also fig. 51). 0. luteus 0. multiporis (138) Wide-headed (width ofhead reaching (149) Neuromasts, often found in group- more than 40 percent of SL; table 9), with a ings oftwo or three, extend along the median relatively long head, reaching nearly 40 per- dorsal ridge from top of the head to base of cent of SL (table 8). Some individuals are the dorsal fin. Scales in, and above and below, known which have heads wider than long. the lateral series with multiple neuromasts. 0. rotundipinnis 0. ctenolepis (139) Pectoral fins nearly perfectly round- (150) All but the scales of the lateral shield ed, rather than with elongate middle rays (fig. are ctenoid in males, with up to 10 ctenii per 41). scale. 0. farfani 0. frontosus (140) An elongate species ofthe luteus group (151) Snout blunt and head rounded (fig. with a relatively straight ventral profile, and 47) throughout all observed life stages. 160 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

0. tschudii green dorsally with an iridescent background (152) There is an increased modal number fading to light yellow-green ventrally. of pectoral fin rays to 19, with an observed 0. polonorum range of from 17 to 19 (table 6). (155) Head rectangular in lateral view, with There is an increased modal number (153) a straight dorsal profile and angular "chin" of vertebrae to 34, with an observed range of (fig. 48). from 33 to 35 (table 3). 0. richersoni 0. empyraeus (154) Unique pigmentation (156) Head, especially in larger adults, ta- pattern char- pers toward mouth (fig. 46), with associated acterized by a black lower jaw margin, faint large preorbital distance (ranging to black spot at between 8 jet dorsal margin ofthe oper- and 12 percent of SL, table 12). culum, and a faint to dark black band at base (157) Lips thick and fleshy in largest adults of the pectoral fin rays. Body a deep olive (fig. 46).

KEY TO ORESTIAS SPECIES This key provides an aid to identification of species. It corresponds with the synapomorphy diagrams of figures 26 through 31. Synapomorphic and autapomorphic characters in the key bear the same number as those in the diagrams. 1 A. (28) Epipleural ribs weakly to strongly bifid in adults; (29) lateral ethmoids relatively far apart (fig. 21 B), and with medial extensions that do not meet or pass dorsal to anterior ramus of the parasphenoid; (30) no marked ontogenetic change in color pattern, with adults showing little or no sexual dichromatism; lateral scales not enlarged, not smooth, with or without granulations; dorsal head scales not enlarged, with or without granulations. 2A 2A. (34) Ventral branchial apparatus elongate, basihyal narrow (fig. 15); (35) fifth ceratobranchials narrow, very close together or fused along midline in adults, toothplates with numerous simple teeth; (36) dorsal gill arches reduced in size relative to ventral gill arches; (37) interarcual cartilage a long, thick rod, equal to or greater in length than first epibranchial bone (fig. 10); (38) modal number of anal fin rays 16 or 17 (table 5); orbits do not project dorsally above surface of head, eyes small to moderate; anal fin roughly parallel to vertebral column, fin base not extending beyond primary ventral profile; second pharyngobranchial bone robust, and with large toothplate bearing numerous simple teeth ...... cuvieri complex. 3A 3A. (60) Modal number of pectoral fin rays 19; modal number of scales in lateral series 33, less than 40 scales; caudal fin truncate, caudal peduncle not constricted; premaxilla a narrow bone with pointed ascending process; dentary narrow, not constricted near medial end. Estrecho Tiquina, Lago Pequenio and Bahia de Puno ...... Orestias forgeti Lauzanne 3B. Modal number of pectoral fin rays less than 19; (44) scales in lateral series 40 or more (table 7); (45) caudal fin lunate, caudal peduncle constricted at base of caudal fin; (46) premaxilla with blunt ascending process, bone wide in lateral view, S shape not pronounced; (47) dentary robust, constricted near its medial end (fig. 20B). 4A 4A. (48) Few or no teeth in outer jaws in adults, juveniles with a few, simple teeth; (49) 6 (rarely 5) branchiostegal rays, with just one ray anterior to or in notch on anterior ceratohyal; (50) modal number of pored scales in lateral series 48 or more (table 7); modal number of vertebrae 32 or 37, not with intermediate number of 34 or 35 (table 3); head not large, length less than 30 percent of SL (table 8); preorbital distance not reaching 12 percent of SL (table 12); moderate to large individuals, not more than 200 mm SL. 5A. SA. (56) Modal number of vertebrae 37 (table 3); (57) scales in lateral series modally 57 (table 7); lower jaw not at 90-degree angle to body axis; ventrum and sometimes either side of median dorsal ridge unscaled. Lago Titicaca and the Cuzco Valley......

...... Orestiaspentlandii Valenciennes 5B. Modal number of vertebrae 32 (table 3); scales in lateral series modally 48 (table 7); (58) lower jaw at 90-degree angle to body axis; (59) body fully scaled with very small, thin irregularly distributed scales. Estrecho Tiquina in Lago Pequeiio, Isla Taquile in Bahia de Puno and the Capachica Peninsula ...... Orestias ispi Lauzanne 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 161

4B. (51) Mouth large, enlarged, recurved unicuspid outer jaw teeth (fig. 32); five branchiostegal rays with just one ray anterior to or in notch on anterior ceratohyal; modal number of pored scales in lateral series 41 (table 7); (52) modal number of vertebrae 34 (table 3); (53) head large, reaching nearly 40 percent of SL (table 8); (54) preorbital distance reaching nearly 14 percent of SL (table 12); (55) largest fishes in genus Orestias, adults reaching 220 mm SL. Lago Titicaca ...... Orestias cuvieri Valenciennes 2B. Ventral branchial apparatus not elongate, basihyal usually rectangular (fig. 14); fifth ceratobranchials moderate, close together but not fused in midline, dorsal gill arches not reduced; interarcual cartilage approximately one-half length of epibranchial bone (figs. 11 and 12); modal number of anal fin rays 14 or 15, rarely 16 or 17 (table 5); (31) orbits project dorsally above surface of head, eyes large; (32) anal fin angled anteriorly away from the body axis, fin base projecting slightly beyond primary ventral profile, creating a distinct transition between abdomen and caudal peduncle; (33) second pharyngobranchial reduced to a U-shaped bone, bears small toothplate with few or no teeth (fig. 11). 6A 6A. (39) Base of the dorsal as well as the anal fin projects beyond primary dorsal and ventral profile (figs. 36 to 40), usually covered with small, thin scales; procurrent caudal fin rays do not lie mostly interior to the body profile, and therefore not relatively close to the vertebral column (fig. 18B) ...... mullericomplex 7A. (69) Modal number of pectoral fin rays 15 (table 5); narrow bodied, with depth less than 23 percent of SL (table10); ventral surface and basal portion ofdorsal, anal and pectoral fins unscaled. Huatayata, Lago Pequeino and Bahia de Puno ...... Orestias gracilis, new species 7B. Modal number of pectoral fin rays more than 15 (table 5); (62) relatively deep bodied, depth ofbody 23 percent or more ofSL (table10); (63) body fully scaled, except in species with granulated dorsal scales. 8A 8A. (70) Modal number of scales in lateral series 36 (table 7); (71) largest fishes in mulleri and gilsoni complexes, reaching an SL of 92 mm; (72) modal number of anal fin rays 13 (table 5); depth ofbody usually less than 26 percent of SL (table10); body relatively fusiform, not laterally compressed. Bahia de Moho, Lago Pequeino, Capachica Peninsula, and ?Bahia de Puno ...... Orestias mullerValenciennesi 8B. Modal number of scales in lateral series less than 36 (table 7); relatively small to medium- sized fishes, 79 mm greatest SL attained; modal number of anal fin rays greater than 13 (table 5); (64) depth of body reaching over 26 percent of SL (table10); (65) body laterally compressed. 9A 9A. (73) Dorsal surface of the head and operculum of males with fleshy, tubercle-like projections covering ctenii of underlying scales, basal portion of dorsal, anal and pectoral fins, and either side of dorsal ridge unscaled, all scales of males apparently with ctenii; modal number of dorsal fin rays 12 or 13 (table 4); depth of caudal peduncle less than 15 percent

of SL (table11); pharyngeal teeth pointed. Isla Taquiri, Lago Pequefio......

...... Orestias tutini Tchernavin 9B. Fully scaled; (66) modal number of dorsal fin rays 16 or 17 (table 4); (67) depth of caudal peduncle 15 percent or more of SL (table11); (68) pharyngeal teeth molariform (figs.11 and 16).lOA OA. (74) Fourth epibranchial bone with huge lateral flange (figs.11 and 12), and associated large adductor IV that fills the cupped surface of the bone; (75) pharyngeal teeth consistently broad (fig. 16); no ventral keel; males and females with faintly mottled color pattern. Bahia de Puno and Bahia de Taman, on the Capachica Peninsula......

...... Orestias crawfordi Tchernavin lOB. Fourth epibranchial bone without huge flange, adductor IV not expanded; pharyngeal teeth molariform, but not as broad as in crawfordi (fig. 16); (76) most laterally compressed fishes in genus Orestias, ventral surface of adults nearly keeled; (77) males with four to five medium to dark brown bars on a lighter brown background, females with fainter bars (although all known specimens faded). Bahia de Moho and ?Bolivia...... Orestias incae Garman 6B. Base of dorsal fin does not extend beyond primary dorsal profile; (40) procurrent caudal fin rays lie mostly interior to the body profile, closer to vertebral column (fig. 18A) giving caudal fin a tapered external appearance ...... gilsoni complex lA. (78) Small fishes with overall body shape characterized by a robust head and relatively 10 narrow caudal peduncle (tables and11) creating an abrupt transition between abdomen and 162 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

caudal peduncle; pigmentation pattern either uniform overall or with barred pattern, but never with marbled pattern. 1 2A 12A. (79) Modal number of pectoral fin rays 17 or 18 (table 6); (90) pigmentation pattern characterized by silvery background with from six to nine irregular bars (figs. 63 and 64); sexual maturity reached at over 25.0 mm SL; scales in lateral series more than 30 (table 7); lower jaw not at right angle to body axis. 1 3A 13A. (93) Head relatively wide and greatly flattened dorsally, its width reaching nearly 23 percent of SL (table 9); (94) caudal peduncle relatively narrow, depth usually less than 10 percent of SL (table 11); modal number of dorsal fin rays 14 (table 4); modal number of vertebrae 32 (table 3). Isla Taquiri and off Tiahuanaco, Lago Pequenio......

...... Orestias gilsoni Tchernavin 1 3B. Head not relatively wide or greatly flattened dorsally, width less than 21 percent of SL (table 9); caudal peduncle not extremely narrow, depth not less than 12.5 percent ofSL (table 11); (95) modal number of dorsal fin rays 12 (table 4); (96) modal number of vertebrae 30 (table 3). Isla Taquiri, Lago Pequenio ...... Orestias taquiri Tchernavin 1 2B. Modal number of pectoral fin rays less than 17 (table 6); pigmentation pattern uniformly pale overall; (81) sexual maturity reached at less than 25.0 mm SL; (82) scales in lateral series 30 or less (table 7); lower jaw not at right angle to body axis. 14A 14A. (97) Body unscaled; (98) modal number of vertebrae 30 (table 3); anal fin rays less than 16. Molinopampa and Sucune, Lago Grande ...... Orestias minimus Tchernavin 14B. Body incompletely scaled; modal number of vertebrae greater than 30; (99) number of anal fin rays 16 or 17 (table 5). Bahia de Uruni, Lago Grande ...... Orestias minutus Tchernavin 12C. Modal number of pectoral fin rays less than 17 (table 6); pigmentation pattern with pale yellow background and dark melanophores overall; sexual maturity reached at over 25.0 mm SL; scales in a lateral series more than 30 (table 7); (83) lower jaw at abrupt (nearly 90-degree) angle to body axis. 1 5A 15A. (100) Modal number of scales in lateral series 37 (table 7); (101) distinctive body shape characterized by relatively narrow (table 9) and deep (table 10) head, with an even more abrupt transition between abdomen and caudal peduncle than diagnostic ofgroup comprising gilsoni, taquiri, minimus, minutus, tomcooni, and mooni; ventrum and basal portion of pectoral, dorsal and anal fin unscaled; modal number ofvertebrae less than 32 (table 3). Near villages of Huatayata and Vilurcuni, Lago Pequefio ...... Orestias tomcooni, new species 15B. Modal number of scales in lateral series 32 (table 7); transition between abdomen and caudal peduncle not as great as in tomcooni; (102) body fully scaled; (103) modal number of vertebrae 32 or 33 (table 3). Bahia de Coata and Bahia de Taman, Lago Titicaca ...... Orestiasmooni Tchernavin 11 B. Small to moderately sized fishes, with overall a relatively robust body and no abrupt transition between abdomen and caudal peduncle; (84) pigmentation pattern characterized by light cream to yellow background covered with dark melanophores arranged in a distinctive marbled pattern (figs. 66, 69, 70 and 72). 16A 16A. (86) Head relatively compressed laterally, width as low as 13 percent of SL (table 9); depth of caudal peduncle less than 12.5 percent of SL (table 11). Estrecho Tiquina, Lago Pe- queio ..... Orestias tchernavini Lauzanne 16B. Head not laterally compressed, its width 16 percent or more of SL (table 9); (85) robust body, depth of caudal peduncle reaching nearly 15 percent of SL (table 11). 1 7A 17A. (87) Lips thick and fleshy (fig. 70); (88) modal number of dorsal fin rays 11 (table 4); basal portion of pectoral fin exposed, not obscured by operculum; anal pouch and papilla cover base of first several anal fin rays; width of head less than 21 percent of SL (table 9); dorsal and ventral profile relatively straight, not arched gently. Bahia de Puno and Lago Pe- queno .. . Orestias imarpe, new species 1 7B. Lips thin, not thick and fleshy; modal number of dorsal fin rays 14 (table 4); (89) basal portion of pectoral fin meets posterior extent of operculum which obscures pectoral fin base; (90) anal pouch and papilla reduced or absent; width ofhead less than 21 percent of SL (table 9); dorsal and ventral profile relatively straight, not arched gently. Bahia de Uruni, Lago Grande .. . Orestias uruni Tchernavin 17C. Lips thin, not thick and fleshy; modal number ofdorsal fin rays 13 or 14 (table 4); basal 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 163

portion of pectoral fin exposed, not obscured by operculum; anal pouch and papilla cover base of first several anal fin rays; (91) body very robust overall, width of head reaches nearly 23 percent of SL (table 9); (92) gentle rise in dorsal profile from tip of snout to base of dorsal fin, and gentle downward slope in ventral profile from tip of lower jaw to base of anal fin (fig. 72). Bahia de Puno, and the Capachica Peninsula ..... Orestias robustus, new species 1 B. Epipleural ribs simple, not bifid; lateral ethmoids relatively close together (fig. 21A), and with medial extensions that usually meet or pass dorsal to the anterior ramus of the parasphenoid; except in one case (mundus), there is a marked ontogenetic change in color pattern, with juveniles and subadult females exhibiting a mottled pattern with a blotched median line and blotched dorsal and lateral surfaces and fins; adult females and subadult to adult males with near uniformly dark dorsal and lateral surfaces, light ventrum and relatively clear fins; (41) lateral scales, from operculum to a point at about the posterior extent of pectoral fin, large and smooth or granulated, forming a so-called lateral shield; (43) dorsal head scales; especially those of median dorsal ridge, enlarged and smooth or granulated, scales of median dorsal ridge large relative to those lateral to the ridge ...... agassii complex 18A. (104) Lateral scales of the lateral shield larger than surrounding scales, overlapping to a great degree, such that only posterior margins of scales are visible; ventrum and basal portion of pectoral, dorsal and anal fins unscaled, otherwise scaled; anterior ribs not large and swollen, body not wide; scales of median dorsal ridge not smooth or granulated. 19A 1 9A. (131) Juveniles through adults all uniformly straw-colored to light brown overall, with a thin, median dark line on lateral surface; modal number of dorsal fin rays 13 (table 4). Cuzco, Peru ...... Orestias mundus, new species 19B. Individuals undergo ontogenetic change in pigmentation pattern such that juveniles have mottled pattern and adults are more uniformly colored; (105) modal number of dorsal fin rays 10 or 11. 20A 20A. (128) Modal number of anal fin rays 11 (table 5); modal number of scales in lateral series 33 (table 7); ventral surface and basal portion of pectoral, dorsal and anal fins unscaled. Lago Ututo, Peru...... Orestias ututo, new species 20B. Modal number of anal fin rays 12 (table 4); (129) when lateral series of scales complete, range 36 to 38 (table 7); (130) squamation often incomplete with dorsal surface sometimes, and ventral surface always, unscaled. Lagos Yanacocha and Llacsha, Peru......

...... O estiasr hardini, new species 1 8B. (106) Body incompletely scaled in particular pattern: scales run posteriorly from the posttemporal in one or two uneven rows, number of rows increasing posteriorly to become a series of four to five rows at base of caudal fin; dorsal scales may be irregularly and sparsely distributed; anterior ribs not large and swollen, body not wide; scales of median dorsal ridge with few distinct concentric striae, no granulations. 21A 21A. (107) Body relatively slender and elongate, depth of body as low as 20 percent of SL (table 10); no sexual dimorphism in scale pattern or number. 22A 22A. (132) Body extremely narrow and elongate, nearly fusiform in largest adults, with narrow head, width nearly as low as 10 percent of SL (table 9); modal number of scales in lateral series 34 (table 7). Pasco Province, Peru...... Orestias gymnotus, new species 22B. Body narrow and elongate, but not nearly as fusiform as in gymnotus, width of head not less than 13 percent of SL (table 9); (133) modal number of scales in lateral series 35 or 36. Rio Santa Eulalia, Peru...... Orestias elegans Garman 21 B. Body not slender and elongate, not fusiform; (108) sexually dimorphic in that females tend to be more fully scaled than males. 23A 23A. (134) Robust species, with relatively long head, reaching nearly 35 percent of SL (table 8); males and females with convex dorsal profile; diploid number of chromosomes not known. Lago Ascot'an, Chile...... Orestias ascotanensis, new species 23B. Head not relatively long, not reaching 32.5 percent of SL (table 8); (135) males and females have a straight dorsal profile, head greatly flattened; diploid number of chromosomes 48 in males and females. Bofedales de Parinacota, Chile...... Orestias parinacotensis Arratia 23C. Head not relatively long, not reaching 32.5 percent of SL; males and females with convex dorsal profile; (136) sexually dimorphic chromosome number, males with diploid chromosome number of 52, females with diploid chromosome number of 50 or 51. Rio Lauca, Parinacota, Chile...... Orestias laucaensis Arratia 1 8C. ( 10) Dorsal head scales as well as lateral scales enlarged and thickened, with few or no concentric 164 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

striae, not overlapping in larger individuals and usually interdigitating along their perimeters, scales cannot be removed easily, giving these fishes an armored appearance, lateral scales with scattered flecks of dark pigment; (109) anterior ribs enlarged and swollen in wide-bodied, large adults. 24A 24A. (1 11) Pigmentation pattern characterized by dark band at base of dorsal, anal and pectoral fin rays, in which band is most prominent; neuromasts in and above and below lateral series single, not in groupings of two or more; body not relatively deep, depth typically less than 29 percent of SL (table 10). 25A 25A. (150) All but scales of lateral shield ctenoid in males, with up to 10 ctenii per scale; modal number of dorsal fin rays 14 (table 4); median dorsal ridge and scales on either side of ridge present and ctenoid in males. Rio Zapatilla, Peru ...... Orestias ctenolepis, new species 25B. Scales of males not extremely ctenoid, if ctenii present, with just a few ctenii per scale only on lateral and posterior scales; (112) modal number of dorsal fin rays 15 or 16 (table 4); median dorsal ridge and scales on either side of ridge present, not ctenoid. 26A. 26A. (151) Snout blunt and head rounded; body fusiform, not laterally compressed. Lago Pequenlo, Bahia de Puno, Capachica Peninsula and Sotalaya ...... Orestias frontosus Cope 26B. Snout slightly tapered, not blunt; (113) body laterally compressed, especially posterior to dorsal fin origin. 27A. 27A. (152) Modal number of pectoral fin rays 19 (table 6); (153) modal number of vertebrae 34 (table 3); body dark brown to black dorsally, pale yellow ventrally, no spot at dorsal margin of operculum. Bahia de Puno, Capachica Peninsula, Guaqui and Taraco, Lago Pequeio ...... Orestias tschudii Castelnau 27B. Modal number of pectoral fin rays 18 (table 6); modal number of vertebrae 32 (table 3); (154) pigmentation pattern characterized by black lower jaw margin, faint to jet black spot at dorsal margin of operculum, and faint to dark black band at base of pectoral fin rays, body deep olive green dorsally with iridescent background, and light yellow-green ventrally. Rio Coata, Peru ...... Orestias richersoni, new species 25C. Scales of males not extremely ctenoid, if ctenii present, there are just a few ctenii per scale only on lateral and posterior scales; modal number of dorsal fin rays less than 15 (table 4); (1 14) median dorsal ridge extremely irregular and incomplete, formed by small, smooth scales. 28A. 28A. (155) Head rectangular in lateral and dorsal view, with straight dorsal profile and angular "chin"; lips relatively thin, not thick and fleshy. Lago Junin, Peru ...... Orestias polonorum Tchernavin 28B. (156) Head in lateral profile, especially in larger adults, tapers toward mouth, with associated large preorbital distance ranging between 8 and 12 percent of SL (table 12); (157) lips thick and fleshy in largest adults. Lago Junin and its tributary streams, Rio Mantaro and Rio Huallaga ...... Orestias empyraeus Allen 24B. Pigmentation pattern without dark band at base of dorsal, anal and pectoral fin rays, body in largest adults dark brown dorsally and laterally, light yellow ventrally, juveniles with mottled pattern; (149) neuromasts, often found in groupings of two or three, extend along median dorsal ridge from top ofhead to base ofdorsal fin, scales in and above and below lateral series with multiple neuromasts. Rio de Lampa, northwestern arm of Rio Coata, Peru ...... Orestias multiporis, new species 24C. Pigmentation pattern without dark band at base of dorsal, anal and pectoral fin rays, body in largest adults dark brown dorsally and laterally, light yellow ventrally, juveniles with mottled pattern; neuromasts in and above and below lateral series single, not in groups of two or more; (1 15) in largest adults, depth of body reaches nearly 35 percent of SL, and is typically more than 29 percent (table 10). 29A 29A. (148) Head squamation irregular, usually with no head scales anterior to the orbit, or on either side of dorsal ridge in young individuals; width of head on average less than 19 percent of SL (table 9). Throughout Urubamba and Titicaca Basins, Salar de Uyuni and northern Chile ...... Orestias agassii Valenciennes 29B. Head fully scaled with large, thickened scales, either smooth or granulated; (1 16) width of head on average 19 percent or more of SL (Table 9). 30A 30A. (146) Lower jaw at 90-degree angle to body axis; (147) dorsal profile convex, rising very sharply from tip of snout to middle of median dorsal ridge; modal number of pectoral fin rays less than 19 (table 6). Rio Guasacona, Lago Urcos and Lago Chinchero, Urubamba Basin ...... Or estiasjussiei Valenciennes 30B. Lower jaw not at abrupt angle to body axis; dorsal profile not convex; (1 17) pectoral fins large, modal number of fin rays 20 (table 6). 3 1A 31A. (145) Dorsal profile nearly straight from tip of snout to anterior base of dorsal fin; body 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 165

not relatively deep, reaching a depth just under 35 percent of SL (table 10); head not relatively wide, less than 23 percent of SL (table 9). Lago Umayo ...... Orestias silustani Allen 31 B. (118) Dorsal profile arched abruptly just posterior to head; (119) body relatively deep, reaching a depth just over 40 percent of SL; (120) head relatively wide, over 28 percent SL (table 9). 32A 32A. (137) Caudal peduncle deep, reaching depth of over 22 percent of SL (table 11); all scales smooth, no granulations. Bahia de Puno, Capachica Peninsula, Lago Umayo ...... Orestiaspuni Tche navin 32B. Depth of caudal peduncle not so greatas in puni, on the average less than 20 percent of SL (table 11); (121) scales from snout to base of dorsal fin, preopercular and opercular regions, and dorsal to pectoral fin thick and covered with dense granulations. 33A 33A. (144) Orbits oriented dorsolaterally, such that in dorsal view, both orbits nearly completely visible; head and scales around and on operculum not swollen. Lago Umayo, Bahia de Puno, and ?Bolivia ...... Orestias olivaceus Garman 33B. Orbits not oriented dorsolaterally; (122) head swollen at a point just dorsal to operculum, scales extremely large and thick, giving body a very angular appearance. 34A 34A. (141) Lower jaw robust, creating prominent "chin"; (142) head relatively long, over 35 percent of SL (table 8); (143) dorsal profile slopes very gently, with no abrupt transition from anterior base of dorsal fin to base of caudal fin, the rays of which flare out slightly at its base; head width less than 29 percent of SL (table 9); scales on operculum not swollen, operculum not arched away from head. Bahia de Puno, Ca- pachica Peninsula, Moho and Huancane ...... Orestias albus Valenciennes 34B. Lowerjaw not robust, no prominent "chin," head length usually under 35 percent of SL, although longer in large adults of luteus (table 8); dorsal profile with abrupt transition between anterior base of dorsal fin and caudal peduncle; (123) head wide, reaching width of over 34 percent of SL (table 9); (124) scales on operculum thick and granulated such that operculum appears swollen and arched slightly away from head. 35A 35A. (140) Elongate species of luteus group with relatively straight ventral profile, and correspondingly a body depth less than 35 percent of SL (table 10); head width less than 35 percent of SL (table 9); fifth ceratobranchials not as close together as in rotundipinnis and luteus, teeth not molariform. Lago Pequeiio and Chililaya, Bolivia ...... Orestiasfarfani, new species 35B. Robust and rotund species of the luteus group; (125) head wide reaching over 37 percent of SL (table 9); (126) pharyngeal teeth molariform with large, rounded cusps; (127) fifth ceratobranchials very close together. 36A 36A. (138) The most wide-headed species in Orestias, width of head reaching over 40 percent of SL, head also relatively long, reaching nearly 40 percent of SL (tables 8 and 9); pectoral fins with elongate middle rays. Localities throughout Lago Titicaca, Lago Umayo, Laguna de Arapa, and Huancane......

...... Orestias luteus Valenciennes 36B. Head not as wide, on the average as in luteus, reaching just under 38 percent of SL (table 9); (139) pectoral fins nearly perfectly rounded, without elongate middle rays. Bahia de Puno, Capachica Peninsula, Lago Pequeno ......

...... Orestias rotundipinnis, new species SYSTEMATIC ACCOUNTS GENUS ORESTIAS4 VALENCIENNES Valenciennes, 1846, by subsequent designation Orestias Valenciennes, 1839, p. 118 (type species of Eigenmann, 1910, p. 461). Orestias cuvieri Valenciennes, in Cuvier and DIAGNOSIS: Defined as a monophyletic group by the following seven synapomor- 4Allen, in Eigenmann and Allen (I1942) used the name phies: no pelvic fins or fin girdles; no vomer; Protorestias (p. 353) to refer to a hypothetical ancestor no first postcleithrum; cartilaginous middle of the genus Orestias; no type species was designated. anal and middle dorsal radials; anterior and The name is still available, and is therefore not listed in posterior bony ceratohyals separated ven- formal synonymy of Orestias. trally by a large gap filled with cartilage; a 166 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 unique squamation and head pore pattern pectoral girdle lowset, first postcleithrum ab- characterized by a dorsal median ridge of sent; posttemporal with ligamentous ventral scales from the head to the dorsal fin and a limb; posttemporal and supracleithrum sep- lyre-shaped arrangement of neuromasts on arate. the dorsal surface of the head; and, angu- Vomer absent; medial ramus of maxilla loarticular lacks a ventral extension parallel straight with pronounced dorsal process with to the retroarticular. or without a groove, dorsal processes nearly DESCRIPTION: First pleural rib on par- meet in the midline, ventral processes nar- apophysis ofsecond vertebra; hypurals fused row; rostral cartilage absent; outer ramus of into a hypural fan; epipleural ribs bifid or not; maxilla robust. transverse processes of vertebrae elongate or Premaxillary ascending processes narrow reduced and cup-shaped. and blunt or pointed, not meeting in the mid- Anal fin inclinators large, nearly meeting line; outer ramus of premaxilla with alveolar or ventral to the division between hypaxial process, indented posteriorly to form an S and epaxial musculature; first proximal ra- shaped ramus. No ethmomaxillary ligament. dial present; middle dorsal and middle anal Uni- or slightly bicuspid outer teeth in one radials cartilaginous. or two irregular rows or outer teeth absent. One dorsal fin ray articulates with first two Dentary with a robust medial extension; dorsal radials; dorsal fin origin opposite or coronoid process on dentary not overlapping slightly in advance of or behind anal fin or- that of anguloarticular; retroarticular not igin. elongate; autopalatine with head angled an- Autopterotic fossa ofmoderate size; lateral teriorly, ventral process elongate, reaching ethmoid enlarged medially or not, when en- quadrate; metapterygoid absent; anguloartic- larged reaching anterior ramus of parasphe- ular lacks a ventral extension parallel to the noid; parasphenoid not broad anteriorly; su- retroarticular. praoccipital and epiotic processes present or Orbital rim free; anterior naris not tubular; absent; nasals flat and broad, nearly meeting cephalic sensory pore system represented by in midline. a lyre-shape pattern of minute neuromasts. Neural arches of the first vertebra open, Ventral surface scaled or not; basal portion not forming a spine, or just meeting to form of dorsal and anal fins scaled or not; scales a blunt spine; first vertebra articulates with often absent from either side of median dor- skull via basioccipital and exoccipital con- sal ridge; scales often large and granulate or dyles; exoccipital condyles reduced; supra- smooth. occipital excluded from formation of fora- Females larger than males, or sexes of ap- men magnum; parietals absent. proximately the same size; dorsal and anal Mesethmoid ossified; pelvic fins and gir- fins rounded, never with fin extensions; cau- dles absent; interarcual cartilage approxi- dal fin rounded, truncate or lunate; pectoral mately one halfof, or equal or greater in length fin rounded or with extended central rays. to, first epibranchial, attaches laterally to sec- REMARKS: The diagnosis and description ond pharyngobranchial with a bony flange; of the genus Orestias are presented in a for- basihyal long and narrow or long and broad mat similar to that ofParenti (198 1) for other anteriorly; no tooth patches on hypobran- cyprinodontiform genera. Diagnoses of chials; no teeth on fourth ceratobranchials; species contain primarily the derived char- dorsal hypohyal absent; no anterior exten- acters whereas descriptions of new species sion ofanterior ceratohyal ventral to the ven- include additional characters that are consid- tral hypohyal; uncinate process on fourth epi- ered to be useful in identification. branchial articulates with that of third; first SPECIES COMPOSITION AND RANGE: Forty- epibranchial narrow or wide at its base; in- three species in the high-altitude lakes and terhyal not ossified; two ossified basibran- tributary streams of the Peruvian, Chilean, chials. and Bolivian Andes from Ancash Province, Preorbital bone flat and broad, sensory ca- northern Peru to Antofagasta Province, nal reduced; dermosphenotic reduced or ab- northern Chile (fig. 1). sent; preopercle with reduced sensory canal; NOTE ON SYNONYMIES: Compilations of, 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 167

FIG. 32. Orestias cuvieri Valenciennes, BMNH 1944.6.6:15-18, female, 180 mm SL. and references to, Orestias species have ap- in Gunther (1866), Garman (1876, Orestias Humboldti. Valenciennes, in Cuvier and peared Valenciennes, 1846, pl. 534, pp. 233-235 (orig- 1895), Eigenmann and Eigenmann (1892), inal description, Lago Titicaca); Terrazas, 1970, Eigenmann (1894a, 1894b, 1910), Cope p. 33 (Bolivia, unconfirmed record). (1876, 1878), Pellegrin (1904a, 1904b, 1906), Orestias Cuvierii. Valenciennes, in Cuvier and Allen, in Eigenmann and Allen (1942), Fow- Valenciennes, 1846, pl. 532. ler (1916), Evermann and Radcliffe (1909, Orestias cuvieri. Tchernavin, 1944a, pp. 153-158 1917), Rendahl (1937) and Zufiiga (1941). (characters, synonymy, distribution); Villwock, These and other references were included in 1962, p. 615 (distribution, ecology); Kosswig the comprehensive synonymies of Tchema- and Villwock, 1964, p. 98 (distribution); Vill- vin (1944a), and for that reason are not re- wock, 1964, pp. 285-291 (discussion of hy- peated here in exhaustive species synony- bridization with Orestias pentlandii in Lago mies. The abbreviated synonymies of Titicaca); Terrazas, 1970, p. 33 (Bolivia, un- previously described Orestias species sum- confirmed record). marize information on synonyms of recog- DIAGNOSIS: An incompletely scaled mem- nized species and orthography of trivial ber of the cuvieri complex most closely re- names, and include, for species described be- lated to the sister species, ispi and pentlandii, fore 1944, Tchernavin's (1944a) reference to distinguished by five derived characters: the species. Also, anatomical references and species with the largest individuals in Ores- distribution records published since 1944 are tias, reaching 220 mm SL; mouth extremely included. large, lowerjaw robust, outer teeth large, uni- To the extent that it was possible to deter- cuspid, recurved and in one large outer row, mine on what material earlier records were and a smaller, irregular inner row; head long, based, identifications were reconfirmed. Rec- its length reaching nearly 40 percent of SL ords for Bolivian species by Terrazas (1970) (table 12); increased modal number of ver- are believed to be based on literature reports tebrae to 34 with an observed range of from and are all unconfirmed. 33 to 35 (table 3); and, preorbital distance greater than in all other Orestias, reaching Orestias cuvieri Valenciennes nearly 14 percent of SL (table 12). Figures 3G, H, I, 32 REMARKS: Meristic and morphometric data Orestias Cuvieri. Valenciennes, 1839, p. 118 (no- for cuvieri are summarized in tables 3 through men nudum); Valenciennes, in Cuvier and Va- 12. lenciennes, 1846, pp. 225-230 (original descrip- Orestias cuvieri Valenciennes has not been tion, Lago Titicaca). collected from Lago Titicaca in recent years, Orestias Humboldtii. Valenciennes, 1839, p. 118 and is presumed to be extinct (see Villwock, (nomen nudum). 1964). The validity of the report of 0. hum- 168 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 33. Orestias pentlandii Valenciennes, BMNH 1944.6.6:22-25, female, 156 mm SL. boldti Valenciennes, a synonym of cuvieri, Bahia de Chucuito: FMNH 41121 (1); Ca- from Bolivia (Terrazas, 1970) is in doubt. pachica Peninsula: USNM 133148 (two Allen, in Eigenmann and Allen (1942) used adults, gill arches of one cleared and coun- the name humboldti for one lot determined terstained for bone and cartilage); BMNH as Orestias mulleri Valenciennes (CAS 46174 1944.6.6:12-14 (three adults, 170-180 mm); and USNM 167731, both formerly IU BMNH 1944.6.6:15-18 (five adults, 185-200 16078), whereas Tchernavin (1944a) treated mm); Puno: MCZ 3927 (four adult specimens 0. humboldti as a synonym of 0. cuvieri. to 220 mm). MATERIAL: Holotype (MNHN A.9593), an RANGE: A midwater species known from adult female, 175 mm SL, collected by M. several localities throughout Lago Titicaca, Pentland, ca. 1839, from Lago Titicaca. recorded from Puno, Bahia de Moho, and the ADDITIONAL MATERIAL: Titicaca Basin: Capachica Peninsula (see Tchernavin, 1944a, Lago Titicaca: BMNH 1862.11.15:46-48 p. 158, and fig. 22 for localities), and from (three subadults, 73-87 mm); BMNH Bolivia, presumably from Lago Titicaca, al- 1862.11.15:78 (one subadult, 54 mm); though, as stated above, the identity of the BMNH 1858.11.5:13-15 (three subadults, specimens reported by Terrazas (1970) has 60-83 mm, one cleared and counterstained not been confirmed. for bone and cartilage); BMNH 1890.2.25:56 (one adult female, 180 mm); BMNH 1862.11.15:31 (one adult female, 180 mm); Orestias pentlandii Valenciennes BMNH 1875.6.7:1-2 (two adults, 180-190 Figures 6A, 10, 33 mm); BMNH 1936.5.6:46 (one adult male, Orestias Pentlandii. Valenciennes, 1839, p. 118 180 mm); USNM 53480 (one adult); MNHN (nomen nudum); Valenciennes, in Cuvier and A.4415; MCZ 3931 (four adults); MCZ 3928 Valenciennes, 1846, pp. 230-232 (original de- (27 juveniles, 17.6-33.1 mm, one cleared and scription, Lago Titicaca). counterstained for bone and cartilage); MCZ Orestias Pentlandi. Valenciennes, in Cuvier and 3926 (six adults); MCZ 31112 (one adult); Valenciennes, 1846, pl. 533. MCZ 6461 (one adult); RMNH 2740 (1); Orestias bairdii. Cope, 1876, p. 185 (original de- RMNH 2733 (3); RMNH 16211 (2); SU 9331 scription, Lago Titicaca). (two adults, 185-220 mm); UMMZ 56895 Orestias bairdi. Eigenmann, 1894a, p. 54 (com- (1); syntypes of Orestias humboldti Valen- piled). ciennes: MNHN 2670 (two subadults, 66- Orestias Petlandi. Zufiiga, 1941, pp. 81-86 (mis- 86.5 MNHN 5829 print). mm); (11 juveniles, 25- Orestias pentlandii. Tchernavin, 1944a, pp. 158- 42 mm); MNHN A.9595 (three subadults, 160 (characters, synonymy, distribution); Vill- 58-74 mm); MNHN A.9596 (two subadults, wock, 1962, p. 615 (distribution, ecology); 70-76 mm); MNHN A.9597 (four subadults, Kosswig and Villwock, 1964, p. 98 (distribu- 67.4-90 mm). tion); Villwock, 1964, pp. 285-291 (discussion 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 169

FIG. 34. Orestias ispi Lauzanne, Holotype, MNHN 1981:605, female, 62 mm SL.

of hybridization with Orestias cuvieri in Lago 1944.6.6:22-25 (five adults, 150-160 mm); Titicaca). BMNH 1944.6.6:36-40 (eight adults, 135- 160 mm); BMNH 1944.6.6:41-48 (11 adults, An incompletely scaled mem- DIAGNOSIS: 160-170 mm); Isla Taquile: AMNH 52198 ber of the cuvieri complex, most closely re- to with small conical or (1). lated ispi and like it Urubamba Basin: Cuzco Valley: MCZ 3941 no outer jaw teeth in adults. One ofthe larger Orestias, individuals reaching 200 mm SL; (3). with the highest number of vertebrae in the RANGE: Recorded from Bahia de Puno (fig. 22), Lago Titicaca in the Titicaca Basin, and genus, the modal number increased to 37, from the Cuzco Valley, in the Urubamba Ba- with an observed range offrom 36 to 38 (table sin. 3); highest number ofscales in a lateral series, the modal number increased to 57, with an Orestias ispi Lauzanne observed range of from 51 to 60 (table 7). Figures 15, 20B, 34 REMARKS: Meristic and morphometric data for pentlandii are summarized in tables 3 Orestias ispi. Lauzanne, 1981, pp. 73-79 (original 12. description, Estrecho Tiquina, Lago Pequefio, through Lago Titicaca, Bolivia). MATERIAL: Syntypes (MNHN A.9594), two adults, 163-168 mm SL, collected by M. DIAGNOSIS AND REMARKS: One of the two Pentland, ca. 1839, from Lago Titicaca. smaller, fully scaled species of the cuvieri ADDITIONAL MATERIAL: Titicaca Basin: complex most closely related to the large, in- Lago Titicaca: MNHN A.9598 (one sub- completely scaled pentlandii, distinguished adult); SU 9306 (two adults, gill arches of one from the other species in the complex by very cleared and counterstained for bone and car- thin, small, irregularly distributed scales that tilage); RMNH 8848 (1); RMNH 15481 (1); are more weakly formed than in the other RMNH 16210 (1); USNM 53478 (8); USNM fully scaled cuvieri complex species, forgeti, 163837 (1); USNM 23246 (2); MCZ 31110 and by a lowerjaw that is at a 90-degree angle (1); MCZ 35059 (2); MCZ 3938 (4); MCZ to the main body axis. 6460 (1); MCZ 3939 (3); MCZ 6459 (1); Young as well as adults of ispi may be con- AMNH 1117 (1); BMNH 1965.10.6:12 (one fused with young of its sister species, pent- adult, 160 mm); BMNH 1890.2.25:55 (one landii, but may easily be distinguished by its adult, 200 mm); BMNH 1852.11.15:28-29 lower number of vertebrae (modal number (two adults, 155-170mm); BMNH 1944.6.6: 32, range 31 to 33 as opposed to a modal 21 (one subadult, 75 mm); BMNH 1944.6.6: number of 37 and a range of from 36 to 38 26-29 (seven adults, 150-170 mm); Lago Ti- for pentlandil) and its lower number ofscales ticaca: Bahia de Puno: USNM 133146 (ex. in a lateral series (modal number 48, range BMNH 1944.6.6:40) (one adult); USNM 41 to 53 as opposed to a modal number of 77613 (3); MCZ 3940 (2); FMNH 81629 (5); 57 and a range of from 51 to 60 for pent- FMNH 41109-41110 (2); FMNH 80323 (1); landil). IU 16084 (at CAS) (two adults, 150 mm); Lauzanne (1981) compared ispi to forgeti AMNH 52110 (one adult, 130 mm); BMNH and to mooni of the gilsoni complex because 170 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 35. Orestias forgeti Lauzanne, Holotype, MNHN 1981:603, female, 73 mm SL.

these species are relatively small, elongate and 23), Isla Taquile in Bahia de Puno, and the are fully scaled; however, he also concluded Capachica Peninsula (fig. 22), all Lago Titi- that ispi was more closely related to pentlan- caca. dii than to mooni. Additional meristic and morphometric data Orestias forgeti Lauzanne are summarized in tables 3 through 12. Also, Figure 35 additional characters not compared among all Orestias in this paper are discussed in Lau- Orestiasforgeti. Lauzanne, 1981, pp. 79-84 (orig- zanne (1981). inal description, Estrecho Tiquina, Lago Pe- MATERIAL: Holotype (MNHN 1981:605), queiio, Lago Titicaca, Bolivia). an adult female, 62 mm SL, collected October DIAGNOSIS: One of the two small, fully 15, 1979, from Estrecho Tiquina, Lago Pe- scaled species of the cuvieri complex, with quefno, Lago Titicaca. Paratypes (MNHN irregularly distributed scales on ventrum 1981:606) 40 adult males and females, 46.5 thinner than dorsal and lateral scales, al- to 76.5 mm, taken along with the holotype. though not so thin and as irregularly distrib- ADDITIONAL MATERIAL: Titicaca Basin: uted as in the other fully scaled species, ispi; Lago Titicaca: AMNH 38400 (four adults, the plesiomorphic species of the cuvieri com- 43-62.5 mm); AMNH 38422 (one subadult, plex, distinguished from cuvieri, pentlandii, 28.5 mm). Bahia de Puno: Isla Taquile: and ispi by a relatively large pectoral fin with AMNH 48641 (232 adults and subadults); an increase in the modal number of pectoral BMNH 1982.12.6:51-120 (70 adults and fin rays to 19, range 18 to 20, as opposed to subadults, 46-54 mm, two cleared and coun- a modal number of 17 with a range of 15 to terstained for bone and cartilage); AMNH 20 (table 6) in the three other species. 48643 (73 adults and subadults); USNM REMARKS: Additional meristic and mor- 228147 (200 adults and subadults); USNM phometric data for this species are summa- 228148 (238 adults and subadults); BMNH rized in tables 3 through 12. Also, additional 1982.12.6:37-50 (14 subadults, 39-56 mm); characters not compared here for all Orestias AMNH 52111 (99 adults, 47-60 mm, three are discussed in Lauzanne (1981). Lauzanne cleared and counterstained for bone and car- compared forgeti with ispi because both are tilage). Capachica Peninsula: AMNH 48640 fully scaled and planktivorous, and with mul- (610 adults and subadults); AMNH 48462 leri and agassii because of the similarity in (269 adults and subadults); AMNH 52112 overall body proportions, however, he made (102 adults and subadults, 47-59 mm, two no firm statement concerning the relation- cleared and counterstained for bone and car- ship offorgeti to any other Orestias species. tilage). MATERIAL: Holotype (MNHN 1981:603), RANGE: Known from the type locality, Es- an adult female, 73 mm SL, collected October trecho Tiquina in Lago Pequefno (figs. 22 and 15, 1979, from Estrecho Tiquina, Lago Pe- 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 171

FIG. 36. Orestias mulleri Valenciennes, Syntype, MNHN A.9605, female, 71 mm SL. queno, Lago Titicaca. Paratypes (MNHN in the modal number of scales in a lateral 1981:604), 40 adult males and females, 56 to series to 36, with a range of from 33 to 38 84 mm, taken along with the holotype. (table 7), as opposed to a modal number of ADDITIONAL MATERIAL: Titicaca Basin: 31 or 32, with a range of from 30 to 36 in Lago Titicaca: Bahia de Puno, between Puno other species of the mulleri complex; and, a and Chucuito, near village of Ojjerani: decrease in the modal number ofanal fin rays AMNH 52113 (four adults, 63-74.5 mm, one to 13, with a range of from 12 to 15 (table cleared and counterstained for bone and car- 5), as opposed to a modal number of 15, with tilage); AMNH 48645 (one adult, 66.5 mm); a range offrom 13 to 18 for the other mulleri AMNH 48647 (one adult, 74 mm); AMNH complex species. Additional morphometric 52114 (two adults, 57.5-59 mm). and meristic data for this species are sum- RANGE: Known from the type locality, Es- marized in tables 3 through 12. trecho Tiquina in Lago Pequefio (figs. 22 MATERIAL: Syntypes (MNHN A.9605), two and 23), and near the village of Ojjerani, be- adult females, 67 to 71 mm SL, collected by tween Puno and Chucuito, Bahia de Puno M. Pentland, ca. 1839, from Guaqui [Guai- (fig. 22). chu], Lago Pequefno, Lago Titicaca. ADDITIONAL MATERIAL: Titicaca Basin: Orestias mulleri Valenciennes Lago Titicaca: BMNH 1862.11.15:32, 33, 59, Figures 14, 21B, 36 73 (4); BMNH 1862.11.15:71 (1); BMNH Orestias Mulleri. Valenciennes, in Cuvier and Va- 1858.11.5:10 (1); AMHN 38423 (two adults, lenciennes, 1846, pp. 240-241 (original descrip- 62.5-68 mm); Bahia de Moho: CAS 46174 tion, Guaqui [Guaichu], Lago Pequeiio, Lago (ex. IU 16078) (20 subadult to adult males Titicaca). and females, 43-75 mm, one cleared and Orestias humboldti (not Orestias humboldti Va- counterstained for bone and cartilage); USNM lenciennes). Allen, in Eigenmann and Allen, 167731 (ex. IU 16078) (11); Bahia de Puno: 1942, pp. 355-357, pl. xvii, fig. 2 (misidentifi- village of Ojjerani, about 10 km S of Puno: cation). AMNH 52211 (one adult, 58 mm); AMNH Orestias mulleri. Tchernavin, 1944a, pp. 222-225 52115 (two adults, 56-60 mm); AMNH (characters, synonymy, distribution); Terrazas, 52116 (two adults, 71-76 mm); AMNH 1970, p. 34 (Bolivia, unconfirmed record). 52117 (two adults, 58-59 mm) Capachica DIAGNOSIS: The largest fishes ofthe mulleri Peninsula: AMNH 52118 (45 adult males and complex, reaching 92 mm SL; fully scaled, females, 64-92 mm, two cleared and coun- with irregularly distributed scales covering terstained for bone and cartilage). the ventrum; most closely related to a group RANGE AND REMARKS: Allen, in Eigen- comprising tutini, crawfordi, and incae but mann and Allen (1942), diagnosed and il- distinguished from these more laterally com- lustrated large specimens of 0. agassii Va- pressed, deeper bodied species by an increase lenciennes (IU 17098) as 0. mulleri 172 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 37. Orestias gracilis, new species, Paratype, AMNH 52121, male, 30 mm SL.

Valenciennes, and 0. mulleri (IU 16078) as though no one morphometric trait separates 0. humboldti Valenciennes, a synonym of 0. gracilis from other Orestias species, it can be cuvieri Valenciennes. Orestias mulleri, there- seen that gracilis (fig. 37) is a relatively slen- fore, is not as abundant and widespread as der-bodied (tables 9 through 11) species of indicated by Allen. It is known with certainty the mulleri complex. It differs in just one me- from several localities in Lago Titicaca, in- ristic character, number of pectoral fin rays, cluding Bahia de Moho, Lago Pequefno, and modally 15 (range of from 14 to 15), which the Capachica Peninsula. One specimen is below the modal number of 17 for the ge- (AMNH 52119) from northeast of Isla Es- nus and 19 for the mulleri complex (table 6). teves in Bahia de Puno is tentatively referred As noted in Phylogenetic Analysis, the closest to mulleri; although, it is most likely an un- relative of Orestias, the genus Kosswigich- described species. thys, has a low number of pectoral fin rays A lectotype was not designated from the also. However, gracilis is clearly a member two syntypes because it is not known whether ofa relatively derived, well-defined subgroup they comprise Valenciennes' entire type se- of Orestias (comprising the mulleri and gil- ries. soni complexes); therefore, the low number of pectoral fin rays in gracilis is considered Orestias gracilis, new species to be secondarily derived, and hence, autapo- Figure 37 morphic. DIAGNOSIS: A relatively small, slender- Adults to 40 mm SL. Males and females bodied species of the mulleri complex, with overall straw-colored, with scattered, small a dorsal and anal fin base that project just melanophores that increase in number (hence slightly beyond the primary body profile; body density) dorsal to the midlateral scales; fins overall relatively light straw-colored, with clear to slightly mottled. scattered, small melanophores that are dense Orbits project slightly above dorsal surface on the dorsal surface; fins relatively clear to of head; dorsal surface slightly to abruptly slightly mottled; pectoral fin small at its base, convex posterior to the head. Ventral surface rays elongate, reaching more than halfway and basal portion ofdorsal, anal and pectoral between base and vertical through origin of fins unscaled. Scales thin and delicate. Males anal fin; modal number of 15 pectoral fin rays with breeding tubercles on dorsal and anal (range of from 14 to 15) as opposed to a fins. Caudal fin truncate. Pectoral fin narrow modal number of 17 pectoral fin rays for the and elongate, reaching just over halfway be- genus, and a modal number of 19 (range of tween pectoral fin base and anal fin. Anal from 15 to 20) for other species ofthe mulleri papilla large (see fig. 37), extending beyond complex (table 6). basal portion of anal fin and fin rays. Basal DESCRIPTION AND REMARKS: Additional portion ofdorsal and anal fins project slightly morphometric and meristic data for gracilis beyond primary body profile. are summarized in tables 3 through 12. Al- Outer jaw teeth small, unicuspid, with 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 173

FIG. 38. Orestias crawfordi Tchernavin, AMNH 52125, male, 56 mm SL. pointed tips that barely protrude through the by one specimen (AMNH 52123) from Bahia epithelium covering upper and lower jaws, de Puno on the western side ofLago Titicaca. arranged in one irregular row. ETYMOLOGY: The trivial name gracilis, a There is one specimen among the six not Latin adjective meaning slender, thin, or sim- considered as part of the type material, but ple, referring to the overall elongate and del- nonetheless tentatively referred to gracilis. icate appearance of this species. This specimen (AMNH 52123), an adult male, 35 mm SL, is slightly deeper bodied Orestias crawfordi Tchernavin than the type and other specimens, and has Figures 11; 12; 16; 17; 38 eight (or possibly nine) discrete dark blotches dorsally, rather than scattered, small mela- Orestias crawfordi. Tchernavin, 1 944a, pp. 226- nophores. Pectoral fin rays number 15. The 227 (original description, Bahia de Taman, color pattern and structure of the caudal fin northeastern side of the Capachica Peninsula, (the procurrent rays appear to be slightly Lago Grande, Lago Titicaca). nearer the vertebral column, as they are in DIAGNOSIS: A deep-bodied, laterally com- the gilsoni complex) suggest that this speci- pressed, fully scaled species of the mulleri men may be a member of an undescribed complex, most closely related to incae, and species within the gilsoni complex, like it with an increased number ofdorsal fin MATERIAL: Holotype (AMNH 52120), an rays (table 4), an increased depth of the cau- adult male, 40 mm SL, collected October 2, dal peduncle (table 11), and molariform pha- 1979 by Tom Coon and M. Reed, near the ryngeal teeth; distinguished from incae by two village of Huatayata, from 0 to 30 m from derived characters: larger, molariform pha- the shore at a depth of from 0 to 1 m, on the ryngeal teeth that have rounded (fig. 16), northeastern side of Lago Pequeino, Lago Ti- rather than slightly pointed cusps; and a fourth ticaca. Paratypes (AMNH 52121), three spec- epibranchial bone with a huge posterior flange imens, 26 to 31 mm, taken along with the (figs. 11 and 12) and an associated large ad- holotype (one cleared and counterstained for ductor IV. bone and cartilage). MATERIAL AND REMARKS: Syntypes ADDITIONAL MATERIAL (not part ofthe type (BMNH 1944.6.6:157-158), one adult male, series): Bahia de Puno, village of Ojjerani: 65.7 mm SL, and one adult female, 65 mm, AMNH 52123 (one adult male, 35 mm); Lago collected September 17, 1937 by the Percy Pequefno: AMNH 52122 (five adults, 33.3- Sladen Titicaca Expedition, at a depth offrom 37.7 mm). 36 to 38 m, from Bahia de Taman on the RANGE: Known from the type locality (figs. northeastern side ofthe Capachica Peninsula. 22 and 23) in Lago Peque-no, and tentatively No lectotype is designated because Tcher- 174 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 39. Orestias tutini Tchernavin, Holotype, BMNH 1944.6.6:544, male, 34 mm SL. navin (1944a, p. 226) listed three specimens, dorsal surface ofthe head and the operculum the two listed here, and a third from Pent- of males covered with fleshy, tubercle-like land's collection, obtained by the BMNH projections covering the ctenii of underlying from the MNHN, in his description of craw- scales; and the basal portion of the dorsal, fordi, and only the first two specimens were anal and pectoral fins and along each side of found in the BMNH. Therefore, the complete the median dorsal ridge unscaled. All scales syntype series has not been examined. of males apparently ctenoid, however, all ADDITIONAL MATERIAL: Titicaca Basin: specimens have been partially descaled. The Lago Titicaca: Bahia de Puno, village of species is known only from the type series of Ojjerani, about 10 km S of Puno: BMNH four specimens which are faded and poorly 1982.12.6:25-34 (10 adult males and fe- preserved. Because the species is incomplete- males, 54-74 mm); AMNH 52124 (three ly scaled, Tchernavin (1944a) placed it in his adults, 43-63 mm); AMNH 52125 (20 adult Group III along with other incompletely males and females, 54-64 mm); AMNH scaled species such as gilsoni. However, I place 52126 (21 adult males and females, 59-69 the species in the mulleri complex along with mm, two cleared and counterstained for bone some fully scaled Orestias species because it and cartilage); AMNH 52127 (nine adult has the defining character of the complex, males and females, 59-79 mm); AMNH that is, the bases of the dorsal as well as the 52128 (four adult males and females, 59-68 anal fin project beyond the primary body pro- mm); Escallami: AMNH 52129 (one sub- file. adult, 36 mm). MATERIAL: Holotype (BMNH 1944.6.6: RANGE: Known from Bahia de Puno and 544), an adult male, 34 mm SL, collected July Bahia de Taman, the northeastern side ofthe 30, 1937 by the Percy Sladen Titicaca Ex- Capachica Peninsula, Lago Titicaca (figs. 22 pedition, at a depth offrom 2.7 to 2.8 m, Isla and 23). Taquiri in Lago Pequefno. Paratypes (BMNH 1944.6.6:545-547) three adults, 32 to 35 mm, Orestias tutini Tchernavin taken along with the holotype. Figure 39 RANGE: Known only from the type locality, Orestias tutini. Tchernavin, 1944a, pp. 219-220 Isla Taquiri, an island in the northeastern (original description, Isla Taquiri, Lago Pe- section of Lago Pequefno (figs. 22 and 23). quefio, Lago Titicaca). DIAGNOSIS AND REMARKS: The smallest Orestias incae Garman fishes of the mulleri complex, reaching 35 Figure 40 mm SL, most closely related to the two other Orestias Incae. Garman, 1895, pp. 155-156 (orig- laterally compressed species of the complex inal description, Bahia de Moho, Lago Titica- (crawfordi and incae), distinguished by the ca). 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 175

FIG. 40. Orestias incae Garman, Syntype, MCZ 3948, female, 57 mm SL.

Orestias incae. Tchernavin, 1944a, pp. 225-226 and morphometric data are summarized in (characters, distribution, synonymy); Terrazas, tables 3 through 12. 1970, p. 33 (Bolivia, unconfirmed record). MATERIAL: Syntypes (MCZ 3948), 12 sub- DIAGNOSIS AND REMARKS: A member of adults to adults, 40 to 57 mm SL, (USNM the mulleri complex most closely related to 120281, ex. MCZ 3948) four adults, 49 to 52 crawfordi from which it is distinguished by mm, (BMNH 1939.7.17:2, ex. MCZ 3948) two derived characters: the most laterally one adult female, 52 mm, collected January compressed fishes in the genus Orestias, with 1875, by S. W. Garman, from Bahia de Moho, ventral surface nearly keeled in larger adults; Lago Titicaca. and, males with four to five irregular, me- RANGE: Known only from the type locality, dium to dark brown vertical bars on a light Bahia de Moho (figs. 22 and 23). Terrazas brown background, females with fainter bars. (1970) lists incae from Bolivia, presumably Orestias incae is known with certainty only also from Lago Titicaca, although this is an from the syntype series, 17 subadult to adult unconfirmed record. males and females collected in 1875 by S. W. Orestias luteus Valenciennes Garman from Bahia de Moho on the north- Figures 6B, 7, 8D eastern side of Lago Grande, Lago Titicaca (fig. 22). Tchernavin (1944a, p. 226) stated Orestias luteus. Valenciennes, 1839, p. 118 (no- that men nudum); Valenciennes, in Cuvier and Va- eight specimens described by Garman lenciennes, 1846, pp. 243-244 (original descrip- (1895) were the only ones known. However, tion, Lago Titicaca); Tchernavin, 1944a, pp. Garman (1895, p. 156) stated that dorsal and 204-209 (characters, synonymy, distribution); anal fin rays were counted on only eight spec- Lueken, 1962, pp. 195-198 (karyology); Vill- imens, without giving an indication of the wock, 1962, p. 615 (distribution, ecology); total number of specimens used in his de- Kosswig and Villwock, 1964, p. 96 (distribu- scription. I have located syntypes of incae in tion); Terrazas, 1970, p. 34 (Bolivia, uncon- three collections (MCZ, USNM, BMNH), and firmed record). do not choose one as a lectotype here because Orestias cypho. Fowler, 1916, pp. 427-429 (orig- I do not know whether all syntypes have been inal description, said to have been taken from located. snake collected near La Paz, Bolivia). This is one of the more distinctive species DIAGNOSIS AND REMARKS: A member of within Orestias, and may show more sexual the luteus group ofthe agassii complex, most dichromatism than is commonly found with- closely related to rotundipinnis, differing from in species of the gilsoni and mulleri com- it by having elongate middle rays of the pec- plexes. However, all specimens are faded in toral fin, and defined by being wide-headed alcohol and do not offer the opportunity to (width of head reaching over 40 percent of describe completely this variation. Meristic SL; table 9), with a relatively long head, 176 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 reaching nearly 40 percent ofSL in the largest subadult); Yungayo: CAS 46178 (ex. IU adults (table 8). 16104) (10 juveniles to subadults); Bahia de There is a great deal of variation among Moho: CAS 46176 (ex. IU 16079) (42 ju- individuals of luteus in the shape ofthe head, veniles to adults, 30.5-100 mm); Sotalaya: particularly in the degree of arching of the AMNH 52137 (four adults, 79-85 mm); Bo- dorsal surface. Also, the head may be dis- livia: ANSP 21920, holotype of Orestias cy- torted to a great degree upon preservation pho Fowler (one adult, 125 mm), said to have and during storage. Fowler (1916) based his been taken from a snake collected near La description of Orestias cypho on one speci- Paz; Lago Umayo: BMNH 1944.6.6:452 (one men, claiming that its convex dorsal profile juvenile); FMNH 88762 (193 juveniles to was arched more abruptly than in luteus. This adults); FMNH 77386 (120 juveniles); CAS specimen was reportedly taken from inside a 48259 (13 juveniles to adults); Huancane: snake. Because of the great variety of head USNM 167733 (6), USNM 167732 (19 ju- shapes, I do not support the division of luteus veniles to subadults), CAS 45794 (ex. IU into several species or subspecies on tbe basis 16086) (40juveniles to subadults), CAS 46177 of differences in head shape, nor do I rec- (ex. IU 16090) (10 juveniles to subadults); ognize 0. cypho. However, additional large Laguna de Arapa: FMNH 77553 (12), IU samples of well-preserved specimens may 16109 (at CAS) (three subadults to adults, present the opportunity to recognize more 42.9-77.5 mm). than one taxon. RANGE: Known from several localities Additional morphometric and meristic data throughout Lago Titicaca, including Bahia de for luteus are summarized in tables 3 through Puno, the Capachica Peninsula, Isla Taquile, 12. Bahia de Moho, Lago Pequeino, Sotalaya, MATERIAL: Holotype (MNHN A.9608), an Huancane, and outside the main body of the adult female, 126 mm SL, collected by M. lake in Lago Umayo and Laguna de Arapa Pentland, ca. 1839, from Lago Titicaca. (figs. 22 and 23). ADDITIONAL MATERIAL: Titicaca Basin: Lago Titicaca: AMNH 38424 (one adult, 105 Orestias rotundipinnis, new species mm); AMNH 38428 (one adult, 150 mm); Figure 41 BMNH 1909.6.15:13 (one adult female, 127 mm); USNM 53487 (1); Capachica Penin- DIAGNOSIS: A wide-headed (width ranging sula: BMNH 1944.6.6:425-429 (nine adult between 29 and 38 percent of SL, table 9), males and females, 91-125 mm); BMNH deep-bodied (depth at posterior extent ofhead 1944.6.6:430-433 (four adults, 112-128 ranging between 29 and 38 percent of SL, ta- mm); BMNH 1944.6.6:435-436 (two adults, ble 10) species ofthe luteus group ofthe agas- 72-125 mm); BMNH 1944.6.6:437-439 (five sii complex, most closely related to 0. luteus adults, 1 16-121 mm); BMNH 1944.6.6:440- Valenciennes, but differing from that species 442 (six adults, 75-115 mm); USNM 133147 by having rounded pectoral fins, rather than (ex. BMNH 1944.6.6:444) (1); BMNH having elongate middle rays of the pectoral 1944.6.6:445-447 (three adults, 106-116 fins. mm); AMNH 52130 (seven adult males and DESCRIPTION: Morphometric and meristic females, 82-104 mm, gill arches oftwo cleared data for this species are summarized in tables and counterstained for bone and cartilage); 3 through 12. These data demonstrate that Bahia de Puno: AMNH 52131 (15 adults, rotundipinnis does not differ from luteus, its 83.5-98 mm); AMNH 52132 (two adults, 73- sister species, in most of these characters, 78 mm); AMNH 52134 (one female, 110 although luteus tends to be a relatively wider- mm); Chucuito: FMNH 41112-41114 (three and longer-headed species than rotundipin- adults); Isla Taquile: AMNH 52133 (one nis. Its most striking difference, and the char- adult, 98 mm); Lago Pequenlo: Bahia by Vi- acter by which rotundipinnis is defined, is the lurcuni: AMNH 52135 (one adult, 87 mm); nearly perfectly rounded pectoral fin in both AMNH 52136 (one adult, 97 mm); Sucune: males and females (fig. 41). In all specimens BMNH 1944.6.6:450-451 (two, juvenile and determined as 0. luteus, the pectoral fin is 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 177

FIG. 41. Orestias rotundipinnis, new species, Holotype, AMNH 52138, female, 109 mm SL. distinctly pointed posteriorly, owing to its teus group species, now known as separate elongate middle rays. collections of rotundipinnis. These may be Dorsal profile ofbody arched abruptly pos- recognized as individual taxa when addition- terior to the head; head scales thick and gran- al material is at hand. ulated; scales of and above operculum thick- MATERIAL: Holotype (AMNH 52138), an ened causing these areas to appear swollen. adult female, 109 mm SL, collected Septem- Preorbital region, ventrum, and basal portion ber 2, 1979 by Tom Coon at a depth of from of the dorsal and anal fins unscaled; basal 1 to 2 m, from just NE of Isla Esteves, Bahia portion ofpectoral fin may be fully scaled (as de Puno, Lago Titicaca. Paratype (AMNH on right side of holotype) or unscaled (as on 52139), an adult female, 95 mm, taken along left side of holotype), or irregularly scaled. with the holotype. Males with ctenii on posterior lateral scales. ADDITIONAL MATERIAL: Titicaca Basin: Adults to 115 mm SL. Body grayish brown Lago Titicaca: AMNH 38426 (one adult male, laterally, darker gray dorsally, ventrum a 115 mm); Lago Pequef-no, near Vilurcuni: grayish cream color. All fins with dark mar- AMNH 52140 (one adult male, 85 mm), near gins of rays. Males with breeding tubercles Huatayata: AMNH 52199 (one subadult, 67.5 on dorsal, anal and pectoral fins. mm); Bahia de Puno: AMNH 52141 (one Outerjaw teeth unicuspid, slightly pointed, adult male, 85 mm); Capachica Peninsula: brown tips just protruding beyond epithe- AMNH 52142 (one adult female, 104 mm), lium covering upper and lower jaws; teeth in BMNH 1944.6.6:434 (two adults, 108-115 a single regular or irregular row. mm), BMNH 1944.6.6:443 (four adults, 95- REMARKS: Just two specimens, which are 115 mm). similar in overall shape, coloration and squa- RANGE: Known from several localities in mation, are given type status. The rest of the Bahia de Puno, the Capachica Peninsula, and material referred to rotundipinnis differs from Lago Pequefio, Lago Titicaca (figs. 22 and these two specimens primarily in two char- 23). acters, either in (1) pigmentation (i.e., they ETYMOLOGY: The trivial name rotundipin- are significantly darker), or (2) shape of the nis, a noun in apposition, from the Latin roots head (i.e., they do not have such a steep rise rotundus, an adjective meaning round, and in the dorsal profile posterior to the head). It pinnis, the ablative plural of pinna, a femi- appears, therefore, that the rounded pectoral nine noun meaning wing or feather, in ref- fin, a distinctly derived character, may even- erence to the rounded margin of the pectoral tually be used to define a larger group of lu- fins. 178 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 42. Orestiasfarfani, new species, Holotype, AMNH 52143, female, 106 mm SL.

Orestias farfani, new species queno, Peru, and a second (115.0 mm SL) Figure 42 from Chililaya, Lago Titicaca, Bolivia. None- DIAGNOSIS: A species of the luteus group theless, it is recognized as a distinct species of the agassii complex that in many char- because the large, well-preserved holotype is acteristics, particularly those pertaining to re- distinguished readily from all other species corded morphometric (tables 8 through 12) in the genus. and qualitative characters of body shape, is Head scales, and those of the median dor- intermediate between the robust yet relative- sal ridge and lateral shield large, thick, and ly slender-headed albus and olivaceus and the covered with dense granulations; scales dor- extremely wide-headed luteus and rotundi- sal to operculum thickened, creating an an- pinnis, more closely related to the last two gular shoulder; opercular scales thickened, species, in sharing with them a head width operculum stands away from head (as in fig. that reaches nearly 35 percent or more of SL 8D); scales present on either side of median (table 9), and having the operculum as well dorsal ridge; ventrum and basal portion of as the head swollen owing to the presence of dorsal and anal fins unscaled. Caudal fin trun- greatly thickened and granulated scales. Dis- cate. tinguished from all other species ofthe luteus Head and body medium brown dorsally, group by having a relatively straight ventral fading to light cream ventrally; minute light profile, and therefore, also a body that is not brown melanophores scattered overall (color as deep as that of albus, luteus, and rotundi- pattern for the holotype; second specimen pinnis (table 10). faded in alcohol). DESCRIPTION AND REMARKS: Morphomet- Outer jaw teeth unicuspid, in one irregular ric and meristic data for this species are sum- row, tips light brown, barely protruding marized in tables 3 through 12. An elongate through epithelium covering upper and lower species within the luteus group, distinguished jaws. by its straight ventral profile; intermediate in MATERIAL: Holotype (AMNH 52143), an many characters between albus and oliva- adult female, 106.0 mm SL, collected August ceus, the relatively slender-headed species, 28, 1979 by Tom Coon and Sr. Acho, at a and luteus and rotundipinnis, the relatively depth of from 1 to 2 m, at a distance of from wide-headed species of the group. 30 to 50 m from the shore, near the village Orestias farfani is known from only two of Vilurcuni, Lago Pequefio, Lago Titicaca. specimens, the holotype (106.0 mm SL), col- ADDITIONAL MATERIAL (no type status): SU lected near the village of Vilurcuni, Lago Pe- 9283, an adult female 115.0 mm, collected 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 179

FIG. 43. Orestias albus Valenciennes, MNHN 05.189, male, 125 mm SL.

by P. 0. Simons (no date recorded), from unique body shape characterized by a dorsal Chililaya, Lago Titicaca, Bolivia. profile of the body that slopes very gently RANGE: Known from the type locality as posteriorly, with no abrupt transition from described above in Lago Pequefno, and Chili- the anterior of the base of the dorsal fin to laya, both Lago Titicaca, in the Titicaca Basin the base of the caudal fin, the rays of which (figs. 22 and 23). flare out slightly at its base (fig. 43). ETYMOLOGY: The trivial name farfani, a Orestias neveui Pellegrin was described genitive, in honor of Sr. L. Edgar Farfan Viz- from one specimen fully scaled on the dorsal cardo, director of the Laboratorio de Puno, surface of the body rather than having scales Instituto del Mar del Peru (IMARPE), to absent from either side of the median dorsal commemorate my appreciation for his gener- ridge, as is the case for the syntypes of 0. osity and support given to research on Lago albus Valenciennes. However, the presence Titicaca. or absence of scales from either side of the median dorsal ridge is variable within nu- Orestias albus Valenciennes merous Orestias species (including, for ex- Figures 8C, 43 ample, cuvieri and pentlandii of the cuvieri complex) and is not considered a character Orestias albus. Valenciennes, 1839, p. 18 (nomen useful for specific differentiation. nudum); Valenciennes, in Cuvier and Valen- Tchemavin (1944a, p. 212) included Pel- ciennes, 1846, pp. 242-243 (original description, Lago Titicaca); Tchernavin, 1944a, pp. legrin's (1906) record of 0. albus from Lago 209-212 (in part, characters, synonymy, distri- Ascotan, Chile in his description. The spec- bution); Terrazas, 1970, p. 34 (Bolivia, uncon- imens on which this record was based firmed record). (MNHN 05.186-188) have been examined Orestias Neveui. Pellegrin, 1904b, pp. 9 5-96 (orig- and determined as 0. ascotanensis, a new inal description, Lago Titicaca). species (pp. 196-197). Orestias neveni. Eigenmann, 1910, p. 461 (mis- Additional meristic and morphometric data print). for 0. albus are summarized in tables 3 Orestias Neveu. Zufniga, 1941, pp. 81-86 (mis- through 12. print). MATERIAL: Syntypes (MNHN A.9607), 2 DIAGNOSIS AND REMARKS: One ofthe larger adults, one male 132 mm SL, one female species ofthe luteus group ofthe agassii com- 142.4 mm SL, collected by M. Pentland, ca. plex, distinguished by three autapomorphies: 1839. a robust lower jaw forming a prominent ADDITIONAL MATERIAL: Titicaca Basin: "chin" (fig. 43); a relatively long head, reach- Lago Titicaca: BMNH 1862.11.15:30 (one ing nearly 40 percent of SL (table 8); and, a adult female, 121 mm); MNHN 05.189 (one 180 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 44. Orestias olivaceus Garman, Syntype, MCZ 3946, male, 85.5 mm SL.

adult male, 125 mm, holotype of 0. neveui synonymy, distribution); Terrazas, 1970, p. 34 Pellegrin); USNM 53522 (two adults); MCZ (Bolivia, unconfirmed record). 27697 (one adult); Bahia de Uruni: BMNH DIAGNOSIS AND REMARKS: A member of 1944.6.6:49 (one adult male 97.8 mm, gill the luteus group of the agassii complex, dis- arches cleared and counterstained for bone tinguished from all other Orestias by orbits and cartilage); Capachica: BMNH 1944.6.6: that are oriented dorsolaterally (see fig. 44), 50 (one adult female, 125 mm); Lago Pe- such that in dorsal view, both orbits are near- quefio, offRio Tiahuanaco: BMNH 1944.6.6: ly completely visible. 51 (one adult male, 110 mm); Bahia de Ta- Tchernavin (1944a) considered 0. silus- man: BMNH 1944.6.6:52 (one juvenile, 44 tani Allen to be, in part, a synonym of oli- mm); Moho: MCZ 3948 (one adult); Bahia vaceus and, in part, a synonym of agassii; de Puno: FMNH 77410 (two adults); MCZ however, see comments for that species be- 3949 (one adult); CAS 40772 (two adults, low forjustification ofits specific recognition. 112-125 mm); AMNH 52197 (one adult fe- In addition, Tchernavin considered 0. affinis male, 100 mm); Chucuito, FMNH 41120 (one Garman to be, in part, a synonym of oliva- adult); village ofOjjerani, S ofPuno: AMNH ceus, and in part, a synonym of agassii. In 52144 (four subadults to adults, 60.0-90.5 fact, Tchernavin (1944a, pp. 199-204) re- mm); AMNH 52145 (one adult, 115 mm); garded afjinis as representing a hybrid be- AMNH 52146 (three adults, 95-108 mm); tween olivaceus and agassii in Lago Umayo. BMNH 1982.12.6:1 (one adult, 122 mm); Orestias affinis Garman is considered herein Huancane: CAS 46180 (ex. IU 16091) (38 (pp. 182-185) to be a synonym of 0. agassli juveniles, 32-55 mm). Valenciennes, although, the possibility that RANGE: Known from several localities in it represents a hybrid has not, ofcourse, been Lago Titicaca, including Bahia de Puno, the refuted. Capachica Peninsula and two of its north- Orestias olivaceus, like all other species in eastern bays (Taman and Uruni), Moho, and the luteus group, has thick granulated scales Huancane (figs. 22 and 23). on the dorsal surface of the head and body. Specimens examined are uniformly medium Orestias olivaceus Garman brown, although Garman (1895, p. 153) states Figure 44 that the species is olive dorsally, and silvery ventrally, and that individuals "oftwo inches" Orestias olivaceus. Garman, 1895, pp. 152-159 are olivaceous. (original description, Lago Umayo); Tcherna- Morphometric and meristic data are sum- vin, 1944a, pp. 199-204 (in part, characters, marized in tables 3 through 12. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 181

FIG. 45. Orestias silustani Allen, Holotype, CAS 44205, female, 84 mm SL.

Orestias olivaceus is known only from the 204 (in part, characters, synonymy, distribu- syntype series of six specimens from Lago tion). Umayo, and one recently collected specimen DIAGNOSIS AND REMARKS: A relatively ro- from Bahia de Puno, Lago Titicaca. Syntypes bust species of the agassii complex, distin- of olivaceus are present in two collections guished by its nearly straight dorsal profile of (MCZ and BMNH). I do not choose one as the head and body in the largest adult (ho- a lectotype here because Garman did not state lotype). The smallest specimens of silustani the number of specimens on which he based are not readily distinguishable from the his description ofolivaceus and I do not know smallest juveniles within the luteus group of whether all syntypes have been located. the agassii complex. Although all the scales MATERIAL: Syntypes (MCZ 3946), five ju- of the adult holotype are smooth, those of a venile to adult males and females, 35 to 85.5 juvenile (60 mm SL) are granulated. The ju- mm SL, (BMNH 1939.7.17: 1, ex. MCZ 3946) venile paratypes are presumed to be conspe- one adult?, 60 mm, all collected 1875 by S. cific with the holotype solely because oftheir W. Garman from Lago Umayo. being collected together. Adult members of ADDITIONAL MATERIAL: Titicaca Basin: the luteus group have densely granulated Lago Titicaca: Bahia de Puno: AMNH 52163 scales. Whether the granulated scales of the (one adult, 93 mm). juveniles catalogued as silustani represent the RANGE: Known from Lago Umayo and Ba- primitive condition ofgranulation, or are ac- hia de Puno, Lago Titicaca (figs. 22 and 23), tually young of another species (e.g., 0. oli- and reported from Bolivia by Terrazas (1970), vaceus, a luteus group species also found in an unconfirmed record. Lago Umayo), awaits the examination ofad- ditional comparative material with more precise collecting data. At present, silustani is Orestias silustani Allen recognized as a distinct species because the Figure 45 largest adult (fig. 45) is readily distinguished from all other Orestias. This specimen (CAS Orestias silustani. Allen, in Eigenmann and Allen, 44205, ex. IU 16097) was indicated as the 1942, pp. 366-367 (original description, Lago in Umayo); Terrazas, 1970, p. 34 (Bolivia, uncon- type (equals holotype) by Allen, Eigen- firmed record). mann and Allen (1942, pl. xix, fig. 1). All Orestias agassii. Tchernavin, 1944a, p. 162 (in remaining 17 specimens of IU 16097 (now part, possibly error in compilation of synony- CAS 44206) are listed herein as possible para- my). types, although it is not clear which of them Orestias olivaceus. Tchernavin, 1944a, pp. 199- Allen wished to be considered part ofthe type 182 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 series. Allen indicated that the description (p. OrestiasAgassizi Var. Senechali. Pellegrin, 1904b, 366) was to be based on five specimens, 55 p. 94 (original description, as a variety, Lago to 86 mm in total length, catalogued as IU Titicaca). 16097. After removal of the holotype from Orestias Agassizi Var. Crequii. Pellegrin, 1904b, IU 16097, it is impossible to tell on which p. 94 (original description, as a variety, Lago four Allen based his de- Titicaca). remaining specimens Orestias uyunius. Fowler, 1940, pp. 63-64 (orig- scription. inal description, Salar de Uyuni, Bolivia). Tchernavin (1944a) listed silustani in the Orestias agassii owenii. not 0. owenii Valen- synonymies of both agassii (p. 162) and oli- ciennes, Tchernavin, 1944a, pp. 177-182 (char- vaceus (p. 199). However, in his table I, p. acters, distribution, synonymy). 144, Tchernavin lists silustani as a synonym Orestias agassii. Tchernavin, 1944a, pp. 162-177 only of olivaceus. Therefore, I believe that (in part, characters, synonymy, distribution); the listing of silustani in the synonymy of Lueken, 1962, pp. 195-198 (karyology); Vill- agassii should be considered a compilation wock, 1964, p. 285 (discussion ofhybridization error. with Orestias olivaceus Garman). Meristic and morphometric data for this Orestias langui. Tchernavin, 1944a, pp. 197-199 (original description, headwaters of the Rio species are summarized in tables 3 through Urubamba, Langui, Peru). 12. Orestias agassi. Parenti, 1981, p. 526 (listed ma- MATERIAL: Holotype (CAS 44205, ex. IU terial). 16097), an adult female, 86 mm SL, collected January 1919, by W. R. Allen, from Lago DIAGNOSIS AND REMARKS: The most wide- Umayo. ?Paratypes (CAS 44206, ex. IU spread, abundant and variable species with 16097), 17 juvenile to subadult males and Orestias; a member of the agassii complex females, 39 to 66 mm, taken along with the most closely related to a group including 0. holotype. jussiei, silustani, puni, and the luteus group RANGE: Known only from Lago Umayo (see fig. 30), and like them, relatively deep (figs. 22 and 23). bodied, particularly in the largest adults (table 10). Orestias agassii is generally distinguished from other species in this group by Orestias agassii Valenciennes a very irregular head squamation pattern, Figures 4A, 8A, 9, 19, 21A usually with no head scales anterior to the Orestias Agassii. Valenciennes, in Cuvier and Va- orbit, and scales absent from either side of lenciennes, 1846, pp. 238-239 (original descrip- the median dorsal ridge, especially in younger tion, Rio de Corocoro and San Antonio de Es- individuals. quilaches, Peru). As recognized by Tchernavin (1944a, pp. Orestias Agassizii. Valenciennes, in Cuvier and 162-177), 0. agassii included 0. empyraeus Valenciennes, 1846, p. xv. Orestias Agassisii. Valenciennes, in Cuvier and Allen, elegans Garman, tschudii Castelnau, Valenciennes, 1846, pl. 536. and possibly silustani (however, see diagnosis Orestias ortonii. Cope, 1876, p. 186 (original de- and remarks for that species, above), all rec- scription, Lago Titicaca). ognized as distinct species in the current re- Orestias ortoni. Eigenmann, 1894a, p. 54 (com- vision. Orestias uyunius Fowler and 0. lan- piled). gui Tchernavin are placed in the synonymy Orestias afJinis. Garman, 1895, p. 152 (original of agassii. Furthermore, several populations description, Lago Umayo). of agassii of Tchernavin are recognized as Orestias tirapatae. Boulenger, 1902, pp. 153-154 distinct species, such as specimens from Lago (original description, Tirapata, Peru). Ascot'an, Chile, described herein as 0. as- Orestias Agassizi. Pellegrin, 1904b, p. 93 (characters). cotanensis, below. One important result of Orestias Agassizi Var. inornata. Pellegrin, 1904b, these taxonomic changes is that agassii is no p. 93 (original description, as a variety, Lago longer recognized as a species ranging Titicaca and Lago Poopo). throughout the distributional limits of Or- Orestias Agassizi Var. typica. Pellegrin, 1904b, p. estias. It is now defined as a species found 93 (original description, as a variety, Lago Ti- throughout the Urubamba and Titicaca ba- ticaca). sins and in northern Chile (Arratia, 1982). 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 183

Most species within the agassii complex one cleared and counterstained for bone and vary little in number of dorsal and anal fin cartilage); BMNH 1858.11.3:347 (24); rays (tables 4 and 5). Recognition of individ- AMNH 20353 (eight juveniles) (two cleared ual species in the past has been based pri- and counterstained for bone and cartilage); marily on differences in overall body shape. MCZ 31111; MCZ 6462; MCZ 3937; MCZ This criterion is used in the present study to 3932; USNM 53516; USNM 23259 (1); recognize several species (e.g., frontosus); ANSP 6983 (1); USNM 127095 (1); San Pa- however, specimens such as those referred to blo: USNM 167743 (5); CAS 48256 (12); uyunius Fowler and langui Tchemavin fall Chucuito: CAS 48251 (1); USNM 167740 within the typical range of variation for one (2); Bahia de Moho: FMNH 77373 (24); or several populations referred to agassii. FMNH 77364 (31); CAS 40771 (ex. IU Some of the characters used previously to 16087) (100 juveniles); CAS 46165 (ex. IU define species or subspecies now placed in 16088) (100 juveniles); UMMZ 179282 (ex. the synonymy of agassii may be found in the IU 16076) (2); Capachica Peninsula: BMNH future to represent consistent differences 1944.6.6:309-320 (102); BMNH 1944.6.6: among populations which might be consid- 308 (1); BMNH 1944.6.6:301-305 (11); ered additional species within the agassli BMNH 1944.6.6:245-249 (six males); complex. This problem is outside ofthe pres- BMNH 1944.6.6:306 (4); BMNH 1944.6.6: ent revision, however. Morphometric and 331 (3); BMNH 1944.6.6:326-330 (10); meristic data for agassil are summarized in BMNH 1944.6.6:291 (ninejuveniles); BMNH tables 3 through 12. 1944.6.6:322-325 (5); USNM 133143 (1); Several different spellings of the specific AMNH 52147 (39 juveniles to adult males modifier of Orestias agassii Valenciennes ap- and females, 25-77.8 mm, four cleared and pear in Valenciennes, in Cuvier and Valen- counterstained for bone and cartilage); USNM ciennes (1846), as listed in the synonymy, 228142; Bahia de Puno: CAS 46172 (24); above. It is not obvious from the work how FMNH 80321 (31); USNM 228145 (20); Valenciennes wished to form a patronym USNM 228140 (30); AMNH 52148 (76 ju- honoring Agassiz. Therefore, the spelling of venile to adult males and females, 36.7-133 Orestias agassii Valenciennes used here is the mm); AMNH 52149 (13 adult males and fe- one that appears above the original descrip- males, 61-116 mm); BMNH 1982.12.6:2-24 tion. It is also the one used by the majority (23, 70-106 mm); AMNH 52153 (six adults, of modem workers on Orestias (e.g., Tcher- 75-1 10 mm); AMNH 52152 (seven adults, navin, 1944a, Villwock, 1964, and Lauzanne, 60-94 mm); AMNH 52151 (four adults, 60- 1981). 85 mm); AMNH 52154 (one adult, 85 mm); MATERIAL: Syntypes (MNHN A.960 1), two AMNH 52155 (37 juveniles, 13.5-23.1 mm); adults, 77 to 79 mm SL, from Rio de Coro- AMNH 52156 (two adults, 67.7-70 mm); Isla coro; (MNHN A.9602), six subadults to Taquile: AMNH 52157 (two adults, 52-62 adults, 49.5 to 59.7 mm, from San Antonio mm); Lago Pequenio: Huatayata: AMNH de Esquilaches, both Titicaca Basin. 52158 (14 juveniles to adults, 31.9-88 mm); ADDITIONAL MATERIAL: Titicaca Basin: Vilurcuni: AMNH 52150 (two adults, 86.5- Lago Titicaca: Syntypes of Pellegrin's (1904b) 112 mm); AMNH 52159 (six adults, 71-74 varieties of agassii: crequii (MNHN 05.181- mm); AMNH 52160 (63 juveniles to adults, 183), three males, 45 to 58 mm SL; senechali 8.3-69.0 mm); Guaqui: FMNH 77374 (202); (MNHN 05.179-180), two females, 39.7 to CAS 46353 (ex. IU 16099) (17). 69 mm, and (MNHN 05.174-178), five males, Lago Umayo: Syntypes of Orestias affinis 52.4 to 57 mm; typica (MNHN 05.171-173), Garman (MCZ 27698) 14 subadults to adults, three adults, 74.6 to 79 mm; inornata 41.5-62.6 mm. (MNHN 05.166-168) three adults, 69.5 to USNM 120280 (ex. MCZ 27698) (four 80 mm; holotype of Orestias ortonii Cope: subadults to adults, 45-54 mm); Rio Ilave: (ANSP 21577) one adult male, 126 mm. UMMZ 179277 (ex. IU 16119) (3); CAS MCZ 3935 (45 juvenile to adult males and 48250 (ex. IU 16119) (five subadults, 41-78 females, 22.0-62.0 mm); BMNH 1862.11.15: mm); Ilave: CAS 46167 (ex. IU 16116) 34 (10 adult males and females, 50-63 mm, (12): Rio Caminaque below Ilave: UMMZ 184 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

179276 (ex. IU 16118) (15); CAS 42534 (ex. Lago Huaypo: UMMZ 179271 (ex. IU IU 16118) (20). 16067) (20); CAS 46157 (ex. IU 16067) (49). Lago Poopo: Additional syntypes of inor- Rio Ilpa: AMNH 52161 (25 subadults to nata Pellegrin: (MNHN 05.169-170) two fe- adults, 31-57 mm). males, 46-54.3 mm. Rio Ayaviri: AMNH 52162 (28 subadults CAS 46352 (ex. IU 16098) (50); USNM to adults, 20-55 mm). 167734 (ex. IU 16098) (14). Lagunillas: UMMZ 179268 (ex. IU 16122) Rio de Lampa: UMMZ 179278 (ex. IU (18); CAS 46159 (ex. IU 16122) (72); BMNH 16124) (20); CAS 46164 (ex. IU 16124) (46). 1944.6.6:342-349 (nine males, one cleared Lago Saracocha: FMNH 77379 (147); and counterstained for bone and cartilage); USNM 167737 (15); Rio Saracocha: USNM BMNH 1944.6.6:350-354 (seven females, 133150 (2); BMNH 1944.6.6:63-68 (6). one cleared and counterstained for bone and Laguna de Arapa: CAS 46354 (ex. IU cartilage); BMNH 1944.6.6:332-341 (20 16110) (4); BMNH 1944.6.6:225-234 (42 ju- males and females, one cleared and counter- veniles). stained for bone and cartilage). Rio de Az'angaro: UMMZ 179287 (ex. IU Urubamba Basin: Lago Urcos: FMNH 16128) (23). 89062 (189); CAS 46348 (ex. IU 16063) (19); Rio de Molino, near Juli: CAS 48260 (ex. USNM 167729 (ex. IU 16063) (5); Cuzco: IU 16127) (71); UMMZ 179286 (ex. IU Hacienda Urco: FMNH 70368 (48). 16127) (18); Rio de Juli: CAS 46152 (ex. IU Pueblo Urubamba: FMNH 94185 (74); 16106) (76). swamps at upper edge of Urubamba: CAS Huancane: CAS 46351 (ex. IU 16115) (70); 48247 (ex. IU 16068) (48). CAS 46350 (ex. IU 16089) (5). Rio Huatanay: Isenchaca: UMMZ 179281 Rio de Tiahuanaco: UMMZ 179280 (ex. (ex. IU 16071) (6). IU 16126) (11); CAS 46161 (ex. IU 16126) Rio Layo: BMNH 1944.6.6:364 (four ju- (29). veniles); BMNH 1944.6.6:361-362 (two ju- Rio Mauri at Calacota: UMMZ 179269 (ex. veniles); BMNH 1944.6.6:363 (onejuvenile); IU 15239?) (23); CAS 46162 (ex. IU 15239?) BMNH 1944.6.6:53-62 (17 juveniles to sub- (38); Rio Mulato: UMMZ 179270 (ex. IU adults). 15250) (15); FMNH 77388 (106); Rio De- Cailloma: FMNH 70366 (105 juveniles to saguadero at Calacota: UMMZ 179272 (ex. adults); FMNH 70369 (49 juveniles to adults). IU 16093) (21); CAS 40774 (ex. IU 16093) Tirapata: Syntypes of Orestias tirapatae (86); CAS 46168 (ex. IU 16093) (66). Boulenger (BMNH 1902.7.29:129-133) four Pampa de Acora: UMMZ 179274 (ex. IU adult females, one juvenile; Rios de Pucara 16112) (13); CAS 48246 (ex. IU 16112) (80); y Porque: UMMZ 179283 (ex. IU 16092) FMNH 77385 (514). (11); CAS 46158 (ex. IU 16092) (31). Crucero Alto: CAS 46173 (ex. IU 14660?) Rio de Langui: FMNH 77372 (62); CAS (1); CAS 46171 (ex. IU 15243) (3); USNM 46349 (ex. IU 16073) (16); CAS 46153 (ex. 167739 (ex. IU 15243) (2). IU 16074) (18); Laguna Langui: CAS 46154 Rio de Santa Rosa: CAS 48249 (ex. IU (ex. IU 16072) (31); BMNH 1944.6.6:365- 17867) (28). 369 (20); BMNH 1944.6.6:370 (1); Langui Rio de Eucalyptus at Eucalyptus: FMNH Village, Upper Urubamba: Holotype of Or- 77378 (66); UMMZ 179275 (ex. IU 16114) estias langui Tchemavin (BMNH 1944.6.6: (20). 152) one adult female, 62 mm; Paratypes Rio Colorado at Viacha: FMNH 88670 (BMNH 1944.6.6:153-156) four adult males. (165); CAS 48253 (ex. IU 16108) (31); UMMZ Salar de Uyuni: department of Potosi at 179273 (ex. IU 16108) (11). Llica: holotype of Orestias uyunius Fowler Rio Grande de Lipez: CAS 46170 (ex. IU (ANSP 68866) one adult female, 81 mm, 16125) (73). Paratypes (ANSP 68867-68891) 25 juveniles Rio de Chupa: UMMZ 179284 (ex. IU to adults; ANSP 83977 (non-type material) 16111) (6); FMMH 88763 (26); FMNH 77391 (73 juveniles to adults, 23-63 mm). (349). RANGE: The most wide-ranging species of 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 185

FIG. 46. Orestias empyraeus Allen, Paratype, CAS 44196, male, 101 mm SL.

the genus Orestias, found throughout local- now CAS 44195, an adult male, collected by ities in the Urubamba and Titicaca basins, W. R. Allen on September 6, 1918 from Lago as well as the Salar de Uyuni, in southern Chinchaycocha (=L. Junin, fig. 24). Addi- Bolivia, and in northern Chile (see Arratia, tional specimens from this IU lot are para- 1982 for Chilean localities). types: CAS 44196 (10 juveniles to adults); CAS 44197 (6). Additional paratype material Orestias empyraeus Allen at CAS: Peru: Pachaca: CAS 44200 (ex. IU Figure 46 15234) (8); Lago Yanamate: CAS 44198 (ex. Orestias empyraeus. Allen, in Eigenmann and Al- IU 15241) (66); Rio Mantaro at Zigzag: CAS len, 1942, pp. 367-370 (original description, 44201 (ex. IU 15236) (originally 53 spec., 48 Lago Junin, Peru). empyraeus, 5 cf. Trichomycterus sp.); Tilar- Orestias elegans. (Not Orestias elegans Garman). nioc: CAS 44202 (ex. IU 15237) (23); Jauja: Allen, in Eigenmann and Allen, 1942, p. 371 CAS 44199 (ex. IU 15233) (83); Huancayo: (in part). CAS 48248 (ex. IU 15232) (50 juveniles to Orestias agassii. Tchernavin, 1944a, pp. 162-177 adults, 18-94 mm); Lago Junin: CAS 46163 (characters, distribution, synonymy). (ex. IU 15239) (35 juveniles to adults, 14-65 DIAGNOSIS: A species of the agassii com- mm); Cerro de Pasco, at railway to Goyllaris- plex most closely related to polonorum, dis- quisga: CAS 46356 (ex. IU 15248) (54 ju- tinguished from it and other species of the veniles to adults, 15-52 mm); Lago Poco- complex by a head that in dorsal profile ta- bamba, near Cerro de Pasco: CAS 46155 (ex. pers toward the snout, and an associated large IU 15246) (17 juveniles to adults, 15-40 mm). preorbital distance, ranging between 8 and Additional paratypes ofcatalogued IU ma- 12 percent of SL (table 12); and, lips thick terial have been identified as follows from and fleshy in largest adults. Additional mor- the collections of the USNM and UMMZ: phometric and meristic data are summarized Lago Junin: USNM 167741 (ex. IU 15239) in tables 3 through 12. (nine subadults to adults, 47-99 mm); Tilar- MATERIAL AND REMARKS: In the descrip- nioc: USNM 167742 (ex. IU 15237) (five ju- tion of empyraeus, Allen, in Eigenmann and veniles to adults, 33-65 mm); Huancayo: Allen (1942, pp. 367-368) listed 13 IU cat- UMMZ 179288 (ex. IU 15232) (four sub- alogue numbers for referred material. He adults to adults, 43-73.5 mm); Rio Mantaro mentioned (p. 369) "the type, 120 mm. long, at Zigzag: UMMZ 179289 (ex. IU 15236) Lake Chinchaycocha (15238)." This speci- (originally 25 specimens, 24 juveniles to adult men, 124 mm SL, and figured in plate xx of empyraeus, 21-50 mm, one juvenile cf. Tri- Allen, has been identified as the holotype, chomycterus sp.). 186 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 47. Orestias frontosus Cope, above, Syntype, ANSP 21555, male, 152 mm SL; below, BMNH 1944.6.6:79-83, male, 91 mm SL (Syntype of Orestias pequeni Tchernavin).

The identification of additional type ma- 44203 (18 juveniles to adults, 17-93 mm, terial in institutions that received IU material two adults, 50-54 mm cleared and counter- is a possibility when these specimens are again stained for bone and cartilage); FMNH 77545 reviewed; however, the identification as type (53), BMNH 1875.10.5:8 (one adult, 108 material requires caution. Much of the type mm). material listed by Allen may not be identified RANGE: Known primarily from Lago Junin positively, lots may contain primarily catfish in northern Peru (fig. 24) and its tributary and may be catalogued as such, or the same streams, as well as Rio Huallaga and the Rio IU catalogue number may have been given Mantaro. to two or more lots and must be checked with the IU catalogues now at CAS. Orestias frontosus Cope Two specimens were removed from IU Figure 47 15238, not recatalogued, but referred by Al- Orestias frontosus. Cope, 1876, p. 187 (original len to 0. elegans. These specimens, two adults description, Lago Titicaca). from Lago Junin, are bleached and have lost Orestias agassii tschudii. Tchernavin, 1 944a, pp. many of their scales; however, I tentatively 182-185 (in part, characters, synonymy, distri- refer them to empyraeus (CAS 48258). bution). ADDITIONAL MATERIAL: Lago Junin: CAS Orestias agassii pequeni. Tchernavin, 1 944a, pp. 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 187

FIG. 48. Orestias polonorum Tchernavin, Lectotype, BMNH 1944.6.6:223, female, 120 mm SL.

185-186 (original description as an infraspecies, although they are always most prominent in Lago Pequefio, Lago Titicaca). recently collected specimens. Morphometric and meristic data for fron- DIAGNOSIS AND REMARKS: A member of tosus are summarized in tables 3 through 12. the agassii complex, closely related to tschu- MATERIAL: Syntypes (ANSP 21555-6), one dii and richersoni, but differing from them in adult male, 155 mm SL, one adult female, having a very blunt snout and rounded head 141 mm SL, collected by J. Orton, 1873, from throughout life (see fig. 47). The large indi- Lago Titicaca. viduals of frontosus also have a very pro- ADDITIONAL MATERIAL: Titicaca Basin: nounced band along the base of the pectoral Lago Titicaca: USNM 127096 (two adults, fin rays, characteristic of a larger group of 82.9-105 mm); BMNH 1890.2.7:57-58 (two agassii complex species to which frontosus adults); Capachica Peninsula: AMNH 52172 belongs. (five adults, 109-140 mm); AMNH 52173 Tchernavin (1 944a) did not consider 0. (26 adults, 91-135 mm); AMNH 52170 (one frontosus Cope to be distinct from 0. agassii adult, 105 mm); BMNH 1944.6.6:183 (1); tschudii Castelnau, and yet he recognized 0. Bahia de Puno: AMNH 52171 (one adult, agassii pequeni as distinct, naming it as new. 111 mm); village of Ojjerani: AMNH 52169 Both tschudii and pequeni were recognized as (eight subadults to adults, 52-127 mm); So- infraspecies of agassii, a rank Tchernavin talaya: AMNH 52166 (27 adults, 80-108 considered to be (p. 144) "oflesser taxonomic mm); Lago Pequeino: off Isla Taquiri: Syn- significance .. ." than the categories ofspecies types of 0. agassii pequeni Tchernavin or subspecies. (BMNH 1944.6.6:79-83) six adult males and Within the species frontosus as recognized females, 86 to 105 mm, collected July 30, here, several degrees of pigmentation of the 1937, at a depth of from 2.7 to 2.8 m, Percy fins are represented, ranging from clear and Sladen Titicaca Expedition. Lago Peque-no: having almost no melanophores in the syn- village ofVilurcuni: AMNH 52164 (one adult, type series of pequeni (BMNH 1944.6.6:79- 114 mm); AMNH 52165 (two adults, 117- 83), dusky in the syntypes offrontosus (ANSP 125 mm); Huatayata: AMNH 52167 (one 21555-6), to pectoral fins with a very pro- adult, 87 mm); AMNH 52168 (three sub- nounced dark band along the basal portion adults, 75-77 mm). of the rays in specimens collected recently RANGE: Known from several localities from Lago Titicaca. The basal portions ofthe throughout Lago Titicaca, including Lago Pe- dorsal and anal fin rays have a weak to very quefno, Bahia de Puno, the Capachica Pen- strong dark band also. It is unknown to what insula, and Sotalaya, on the eastern shore (figs. degree these pigments fade in preservation, 22 and 23). 188 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 49. Orestias elegans Garman, Syntype, MCZ 27694, female, 44 mm SL.

Orestias polonorum Tchernavin MATERIAL: Lectotype (BMNH 1944.6.6: Figure 48 223), an adult female, 120 mm SL, collected Orestias polonorum. Tchemavin, 1 944a, pp. 195- 1937 by H. Morrison from Lago Junin, des- 197 (original description, Lago Junin, Peru). ignated herein. Paralectotypes (BMNH 1875.10.5:4-7) four adults, 105 to 117 mm, DIAGNOSIS AND REMARKS: A species of the collected by Jelski, member of the Count K. agassii complex, most closely related to em- Branicki Expedition to South America in pyraeus with which it shares an extremely 1866-1867, from Lago Junin. irregular and incomplete median dorsal ridge RANGE: Known only from the type series, formed by small, smooth scales; distin- Lago Junin, Peru (fig. 24). guished from all other species ofthe complex by a head that is rectangular in lateral view, with a straight dorsal profile and angular Orestias elegans Garman "chin" (fig. 48). Figures 18B, 49 The five syntypes, the only known speci- Orestias elegans. Garman, 1895, p. 149 (original mens, are faded in alcohol except for the pres- description, headwaters ofthe Rio Santa Eulalia ence of scattered, discrete dark spots distrib- where it meets the Rio Rimac, Peru). uted randomly over the dorsal and lateral Orestias agassii elegans. Tchernavin, 1944a, pp. surfaces. The largest specimen (BMNH 186-189 (characters, synonymy, distribution). 1944.6.6:223) is figured by Tchernavin DIAGNOSIS AND REMARKS: A slender and (1944a, p. 195) and is therefore chosen as the elongate species ofthe agassii complex, most lectotype. Morphometric and meristic data closely related to gymnotus, another elongate for these five specimens are summarized in species, both members of a larger group of tables 3 through 12. agassii complex species that are defined as a

FIG. 50. Orestiasjussiei Valenciennes, Syntype, MNHN A.9599, female, 80.5 mm SL. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 189 monophyletic group by a reduced squama- bust species ofthe agassii complex with large, tion pattern characterized by a unique dis- thick, smooth head and body scales, distin- tribution of lateral scales (pp. 123-126); ele- guished from other members of the complex gans is distinguished by an increase in the by: a lower jaw at a 90-degree angle to the modal number of scales in a lateral series to body axis (figs. 20A and 50); and, a convex 35 or 36, with an observed range of from 34 dorsal profile, rising very sharply from the tip to 38 (table 7). Its closest relative, gymnotus, of the snout to the middle of the median has a modal number of 34 scales in a lateral dorsal ridge. The latter character, a convex series, with an observed range of from 31 to dorsal profile, is evident in juveniles as well 35 scales (table 7). Additional morphometric as large adults (fig. 50). and meristic data for elegans are summarized Morphometric and meristic data forjussiei in tables 3 through 12. are summarized in tables 3 through 12. Most specimens examined are bleached; Tchernavin (1944a, p. 177) stated that the however, some do have a silvery ventral and types of Orestias owenii Valenciennes, of the lateral surface of the body, as noted by Gar- collection of the Museum National d'His- man (1895, p. 149) in his original description. toire Naturelle, were lost. He based his di- Syntypes have been located in the collections agnosis of Orestias agassii owenii (pp. 177- of the MCZ and BMNH. However, it is not 182) on specimens of Orestias agassii from known whether all syntypes have been ex- Lago Urcos, in the Urubamba Basin, the type amined, and therefore, no lectotype is des- locality of 0. owenii Valenciennes. The types ignated. ofValenciennes' owenii, however, are still in MATERIAL: Syntypes (MCZ 27694) 10 ju- existence in the MNHN. I examined these venile to subadult males and females, 32 to and determined them to be the young of 0. 49 mm SL (BMNH 1939.7.17:3, ex. MCZ jussiei, known also from the Urubamba Ba- 27694) one subadult, 48 mm, collected by S. sin. W. Garman, in small lakes among the head- Orestias jussiei puni Tchernavin is treated waters ofthe Rio Santa Eulalia where it meets as a valid species in the present revision (pp. the Rio Rimac. 189-190). ADDITIONAL MATERIAL: Peru: Rio Santa MATERIAL: Syntypes (MNHN A.9599), Eulalia: CAS 46179 (ex. IU 15242) (46 ju- three adults, 68 to 80 mm SL, collected from venile to adult males and females, 24-87 mm, the Rio Guasacona in the Urubamba Basin. four of the juveniles to adults, 37-64 mm ADDITIONAL MATERIAL: Urubamba Basin: cleared and counterstained for bone and car- Lago Urcos: Syntypes of Orestias owenii Va- tilage). lenciennes (MNHN A.9606), five juveniles, RANGE: Known only from the Rio Santa 33 to 55 mm; Lago Chinchero: CAS 46175 Eulalia in northern Peru (fig. 24), on the Pa- (ex. IU 16066) (37 juvenile to adult males cific slope of the Andes. and females, 30-84 mm, four cleared and counterstained for bone and cartilage); USNM Orestias jussiei Valenciennes 167730 (ex. IU 16066) (7). Figures 13, 20A, 50 RANGE: Known from several localities in Orestias Jussiei. Valenciennes, in Cuvier and Va- the Urubamba basin (Rio Guasacona, Lago lenciennes, 1846, pp. 235-238 (original descrip- Urcos, and Lago Chinchero) (fig. 24). tion, Rio Guasacona, Peru). Orestias Jussieui. Castelnau, 1855, pl. 27, fig. 1. Orestias puni Tchernavin Orestias Owenii. Valenciennes, in Cuvier and Va- Figures 8B, 51 lenciennes, 1846, pp. 241-242 (original description, Lago Urcos, Peru). Orestiasjussieipuni. Tchernavin, 1944a, pp. 194- ?Orestias Oweni. Rowntree, 1903, p. 69 (report of 195 (original description, as a subspecies, Bahia ductus pneumaticus located to the right ofmid- de Puno, Lago Titicaca). dorsal line). DIAGNOSIS AND REMARKS: A robust mem- Orestias jussiei. Tchernavin, 1944a, pp. 189-191 (in part, characters, distribution, ber ofthe agassii complex, distinguished from synonymy). all other species in the complex (minus 0. DIAGNOSIS AND REMARKS: A relatively ro- luteus) by: a very deep caudal peduncle, 190 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 51. Orestias puni Tchernavin, AMNH 52175, female, 111 mm SL. reaching a depth of over 22 percent of SL estias (the others are agassii, ascotanensis, (table 11). Orestias luteus is the only other and laucaensis of the agassii complex) with species in the genus in which individuals have members in Chile, most closely related to the such a deep caudal peduncle. However, puni latter two species. Orestias parinacotensis is is readily distinguished from all species ofthe distinguished by a straight dorsal head profile luteus group by large, thick smooth scales on and a head that is greatly flattened in both the head and body, rather than granulated males and females (see fig. 52). scales which is diagnostic ofthe luteus group. REMARKS: Orestias parinacotensis Arratia Morphometric and meristic data are sum- has not been examined, as notice of deposi- marized in tables 3 through 12. tion of specimens was received just prior to MATERIAL: Syntypes (BMNH 1944.6.6: publication of this revision. The holotype of 179-180), two adults, one male 124 mm SL, this species, as well as that of 0. laucaensis, one female 122 mm, collected August 1, 1937 is deposited in the Coleccion de Ictiologia, (BMNH 1944.6.6:181), one adult, 117 mm, Laboratorio de Biologia, Universidad de collected August 5, 1937 (BMNH 1944.6.6: Chile, Santiago-Sur. Additional specimens 182) one adult, 120 mm, collected July 22, (including paratypes) have been deposited in 1937, all collected by the Percy Sladen Titi- the collection of the University of Kansas, caca Expedition, from Bahia de Puno, Lago Museum of Natural History. Morphometric Titicaca. and meristic data presented in the current ADDITIONAL MATERIAL: Titicaca Basin: paper (tables 4 through 11) are from a pre- Lago Titicaca: AMNH 52174 (one adult, 80 publication copy of Arratia (1982). mm); Capachica Peninsula: AMNH 52175 RANGE: Known from the Chilean Alti- (two adults, 111-120 mm); Lago Umayo: plano, in Parinacota, Chile (fig. 25). See also CAS 46355 (ex. IU 16095) (8). Arratia (1982) for remarks on the locality. RANGE: Known from Bahia de Puno and the Capachica Peninsula in Lago Titicaca, and Lago Umayo (figs. 22 and 23). Orestias laucaensis Arratia Figure 53 Orestias parinacotensis Arratia Orestias laucaensis. Arratia, 1982, pp. 103-104 Figure 52 (original description, Rio Lauca, Parinacota, Orestias parinacotensis. Arratia, 1982, pp. 100- Chile). 103 (original description, Bofedales de Parina- DIAGNOSIS: One ofthe four species (agassii, cota, Chile). parinacotensis, ascotanensis, in addition to DIAGNOSIS: One of the four species of Or- laucaensis) ofOrestias with members in Chile, 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 191

-. ...

FIG. 52. Orestias parinacotensis Arratia, above, Paratype, KU 19213, female, 62.3 mm SL; below, Paratype, KU 19213, male, 44.5 mm SL.

most closely related to the last two species, See also Arratia (1982) for remarks on the all ofthe agassii complex. Orestias laucaensis locality. is the only species of Orestias known to have sexually dimorphic chromosome numbers. Orestias tschudii Castelnau The diploid number of chromosomes is in- Figure 54 creased to 52 in males, and 50 or 51 in fe- Orestias Tschudii. Castelnau, 1855, pp. 51-52 males, as opposed to a plesiomorphic diploid (original description, Lago Titicaca). number for teleosts of 48 in both males and Orestias Tschudi. Bleeker, 1860, p. 487 (com- females (see discussion, above). piled). REMARKS: Orestias laucaensis Arratia has Orestias agassii tschudii. Tchernavin, 1944a, pp. 182-185 (in part, characters, synonymy, distri- not been examined (see remarks under 0. parinacotensis Arratia, above). The holotype bution). of this species is deposited in the Coleccion DIAGNOSIS: A member of the agassui com- de Ictiologia, Laboratorio de Biologia, Uni- plex, with large, thick, smooth head and body versidad de Chile, Santiago-Sur. Additional scales, closely related to richersoni, diagnos- specimens (including paratypes) have been able by: an increased modal number of 19 deposited in the collection of the University pectoral fin rays, with an observed range of ofKansas, Museum ofNatural History. Mor- from 17 to 20 (table 6), and an increased phometric and meristic data for this species modal number of 34 vertebrae, with an ob- (tables 4 through 1 1) are from a pre-publi- served range of from 33 to 35 (table 3). cation copy of Arratia (1982). REMARKS: See remarks under 0. frontosus RANGE: Known from Rio Lauca, Parina- (p. 187) for pertinent comments on this cota, Chile, on the Chilean Altiplano (fig. 25). species. 192 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

it :-r*,r j 14...t.,.rll -1 '"

FIG. 53. Orestias laucaensis Arratia, Paratype, KU 19124, female, 67.4 mm SL.

MATERIAL: Syntypes (MNHN A.9604) name implies, one of the nearly unscaled three adults, 118 to 129 mm SL, collected species in the agassii complex, with few or from Lago Titicaca. no head scales, an incompletely formed me- ADDITIONAL MATERIAL: Titicaca Basin: dian dorsal ridge, and no lateral scales dorsal Lago Titicaca: AMNH 38425 (one adult, 125 or ventral to the lateral, median rows ofscales mm); AMNH 38427 (one adult, 145 mm); (fig. 55). Bahia de Puno: BMNH 1944.6.6:162-167; DESCRIPTION AND REMARKS: Morphomet- BMNH 1944.6.6:168-172; BMNH 1944.6.6: ric data summarized in tables 8 through 12 204-211 (12); BMNH 1944.6.6:212-217; serve to show that gymnotus is a very slender AMNH 52177 (19 adult males and females, and elongate species within the agassii com- 79-102 mm); FMNH 88806 (9); Capachica plex; meristic data summarized in tables 3 Peninsula: BMNH 1982.12.6:124-131 (eight through 7 for the species fall within the av- adults, 99-112 mm); AMNH 52176 (47 erage range ofvalues for the agassii complex. adults, 88-132 mm); BMNH 1944.6.6:190- The squamation pattern of gymnotus 197; BMNH 1944.6.6:218-221 (4); BMNH (found to a certain degree among related 1944.6.6:198-203 (7); BMNH 1944.6.6:184- species, including ascotanensis and elegans) 187 (8); CAS 48252 (16 subadults to adult, is relatively invariant within the species. The 40-131 mm); Huancane: BMNH 1944.6.6: pattern can be seen in the photograph of the 222 (1); Lago Peque-no: Taraco: BMNH holotype (fig. 55), an adult female. Scales run 1944.6.6:94-101 (one cleared and counter- posteriorly from the posttemporal in one or stained for bone and cartilage); BMNH two rows. The number ofscale rows increases 1944.6.6:84-93 (20); Guaqui: BMNH posteriorly to become a series of four to five 1944.6.6:142-151; BMNH 1944.6.6:122- rows at the base of the caudal fin. Scales are 131; BMNH 1944.6.6:132-141 (one cleared absent from the ventrum, and basal portions and counterstained for bone and cartilage); ofthe pectoral, dorsal, and anal fins. Despite BMNH 1944.6.6:112-121 (uveniles). the sparse squamation, it can still be seen (fig. RANGE: Known from several localities in 55) that gymnotus has enlarged, smooth scales Lago Titicaca, including Bahia de Puno, the just posterior to the operculum, diagnostic of Capachica Peninsula, Guaqui and Taraco, the agassii complex. Lago Pequefio (fig. 22). Adults to over 60 mm SL. Color patterns ofjuvenile through adult males and females Orestias gymnotus, new species typical of those described for Pellegrin's Figure 55 (1 904b) varieties of 0. agassii, above. Large DIAGNOSIS: A very slender and elongate adults are medium to light brown overall, species of the agassii complex, nearly fusi- with a pale yellow ventrum; usually, there are form in overall body shape as exemplified by minute dark brown melanophores overall. its relatively narrow head (table 9) and low Juveniles have an incomplete to complete depth ofthe body (table 10), and as the trivial dark brown midlateral line of pigment, and 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 193

FIG. 54. Orestias tschudii Castelnau, Syntype, MNHN A.9604, male, 129 mm SL (posterior section of body partially descaled). large, dark blotches dorsally, and often sev- cality as described above (no type status): CAS eral blotches ventral to the midlateral line. 51310 (3); Pasco Province: Arroyo Tusi at Fins with small brown spots, or clear. Caudal Pocobamba (4154 m): FMNH 77905 (122), fin truncate. CAS 48257 (ex. IU 16061) (73); Arroyo Outer jaw teeth unicuspid, in one irregular Cuchis: CAS 40768 (37). row. Tips of teeth slightly pigmented, barely RANGE: Known from several localities in protruding beyond thick epithelium covering Pasco Province, northern Peru, just north of upper and lower jaws. Lago Junin (fig. 24). MATERIAL: Holotype (CAS 40700, ex. IU ETYMOLOGY: The trivial name gymnotus, 16102), an adult female, 62.0 mm SL, col- an adjective, from the Latinized Greek root lected 1918 by C. H. Eigenmann, from Cha- gymnos, meaning naked or lightly clad, and llhaucocha, near Quishuarcancha, northwest the Latin adjectival suffix -tus denoting pos- of Cerro de Pasco, Pasco Province, northern session, in reference to the sparse squamation Peru, at an elevation of approximately 4062 pattern. m. Paratypes (CAS 51309) 38 juvenile to adult males and females, 15.0 to 52.0 mm, three Orestias hardini, new species cleared and counterstained for cartilage and Figure 56 bone, taken along with the holotype. DIAGNOSIS: A species of the agassii com- ADDITIONAL MATERIAL: From the type lo- plex, most closely related to 0. ututo, another

FIG. 55. Orestias gymnotus, new species, Holotype, CAS 40700, female, 62 mm SL. 194 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 56. Orestias hardini, new species, Holotype, AMNH 52178, male, 29 mm SL. new species (p. 200) with which it shares a yond epithelium covering upper and lower low number ofdorsal fin rays (modally 10 in jaws. ututo; 11 in hardini, observed range in both Orestias hardini is a small species; a gravid of from 10 to 12; table 4). Distinguished by female 38.5 mm SL is the largest known spec- a high number of scales in a lateral series; imen. However, as noted in Phylogenetic when the series is complete, scale number Analysis, the nature and distribution of di- observed to range from 36 to 38 (table 7). minutive body size in Orestias is unknown. Squamation incomplete, with dorsal surface Killifishes apparently may continue to grow of some specimens and ventral of all, devoid at sexual maturity. Large, mature adults oth- ofscales. Lateral scale series often incomplete erwise referable to hardini should not be or absent. placed in another new species until the pos- DESCRIPTION AND REMARKS: Morphomet- sibility that hardini continues to grow at sex- ric and meristic data for this species are sum- ual maturity is investigated further. marized in tables 3 through 12. Orestias har- MATERIAL: Holotype (AMNH 52178, ex. dini is most closely related to 0. ututo with AMNH 38415), an adult male, 29.0 mm SL, which it shares a low number of dorsal fin collected June 5, 1979 by Tim Hardin, from rays (see comments above and table 4). It is Lago Yanacocha, Peru (approx. 1 0°S, most easily distinguished from ututo by a high 77°15'W). Paratypes (AMNH 38415), four number of scales in a lateral series, ranging juvenile to adult males and females, 19.0 to from 36 to 38 in hardini, as opposed to a 38.5 mm, one cleared and counterstained for range of 32 to 37, with a mode of 33 in ututo bone and cartilage, taken along with the ho- (table 7). The squamation of hardini is also lotype; (AMNH 38411), four juveniles, 13.5 often irregular, with the lateral series ofscales to 20.0 mm, collected June 13, 1979, by Tim often incomplete or absent. Ventral surface Hardin, from Lago Llacsha, Peru (approx. and basal portions of pectoral, dorsal, and 9045'S, 77033'W). anal fins unscaled. RANGE: Known from lagos Yanacocha and In other meristic and morphometric char- Llacsha (fig. 24) two small isolated lakes in acters, hardini differs little from the average northern Peru, outside the Titicaca and Uru- values of these data for the agassii complex. bamba basins. The Lago Llacsha locality rep- Color pattern is typical of that described for resents the northernmost limit of the genus. Pellegrin's (1904b) varieties of 0. agassii (pp. ETYMOLOGY: The trivial name hardini, in 131-133); that is, background is cream to honor of Tim Hardin, in appreciation of yellow, with a distinct dark midlateral line, his collecting Orestias in northern Peru. and brown blotches dorsally. Dorsal, pectoral, and caudal fins with small brown mela- Orestias ctenolepis, new species nophores; anal fin clear. Blotches in dorsal Figures 57A, B fin may be arranged in two regular rows. DIAGNOSIS: A slender species ofthe agassii Outer jaw teeth unicuspid, in a single, ir- complex, in which males possess extremely regular row. Tips brown, protrude slightly be- ctenoid scales (with up to 10 ctenii per scale) 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 195

FIG. 57. Orestias ctenolepis, new species, A. Holotype, AMNH 52179, male, 68 mm SL; B. Paratype, AMNH 52180, female, 56.8 mm SL. over most of the body, including the dorsal bercles on the anal, dorsal and pectoral fins. surface of the head and median dorsal ridge; Adults to 68 mm SL. only those scales immediately posterior to the Individuals of ctenolepis apparently go base of, and medial to, the pectoral fin (i.e., through the same ontogenetic changes in col- those of the lateral shield) are strictly cycloid or pattern described for Pellegrin's varieties and have not been observed to possess ctenii. of 0. agassii (pp. 131-133); that is, juveniles All fins of both males and females delicate and subadult to adult females (fig. 57B) have and not covered with a thick membrane; up- a mottled midlateral line that becomes more per and lower jaw margins "frilled," that is pronounced with increased size (i.e., SL), and covered with small, delicate papillae. they are mottled over the dorsal surface of DESCRIPTION AND REMARKS: Morphomet- the head and body immediately ventral to ric and meristic data for ctenolepis are sum- the midlateral line. In adults males and some marized in tables 3 through 12. It is a rela- adult females, both the dorsal surface and the tively slender-bodied species of the agassii midlateral line become darker with increased complex, but not so slender as gymnotus, and size, the midlateral line expanding with the not differing significantly in meristic and result that the largest adults are nearly uni- morphometric data from the average values formly deep chocolate brown dorsally and for these characters in other species of the laterally (fig. 57A). Both males and females complex. Distinguished from all other Or- are light cream ventrally. There is a faint bar estias by the males having nearly all scales at the base of the pectoral fin rays. (except those of the lateral shield) with up to Basal portions of the dorsal, anal and pec- 10 ctenii per scale. The maximum number toral fins, and ventrum unscaled. The holo- of ctenii occurs on the scales posterior to the type and several paratypes (one as small as lateral shield. Males also have breeding tu- 40 mm SL) have the area on either side of 196 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 58. Orestias ascotanensis, new species, Holotype, USNM 236790, female, 54 mm SL (caudal fin broken). the median dorsal ridge unscaled. In some of laucaensis, the two otlher Orestias species en- the smaller specimens, scales have been lost, demic to Chile, sharing with them a sexual making an accurate count ofscales in a lateral dimorphism in which females tend to be more series impossible (see table 7). fully scaled than males; females have several The "frilled" upper and lowerjaw margins complete lateral scale rows, whereas, males help to identify the species, but this state does usually have one anteriorly, gradually in- not represent a unique characteristic ofcteno- creasing to several posteriorly. Head rela- lepis, as such papillae occur in some popu- tively long for a species ofthe agassii complex lations of 0. agassii not recognized as distinct not a member of the luteus group, and the taxonomically in this study. one character by which ascotanensis is readi- Outer jaw teeth unicuspid, with clear, un- ly distinguished; the length ofthe head reach- pigmented tips, arranged in a single, irregular es nearly 35 percent of SL (table 8). row, which could be interpreted as two rows DESCRIPTION AND REMARKS: Morphomet- in some specimens. ric and meristic data for ascotanensis are MATERIAL: Holotype (AMNH 52179), an summarized in tables 3 through 12, showing adult male, 68.0 mm SL, collected September that whereas ascotanensis differs little in me- 7, 1979 by Tom Coon, from the headwaters ristic characters from the other species in the ofthe Rio Zapatilla, in Rio Camellaque, near agassii complex, the morphometric charac- Km 24 on the Ilave-Mazo Cruz Road, near ters indicate that it is a robust species, most Tiutiri, Chucuito Province, Peru, Rio Za- easily distinguished by its relatively long head patilla drainage. Paratypes (AMNH 52180), (table 8). eight juvenile to adult males and females, Sexual dimorphism and dichromatism 37.0 to 63.0 mm, taken with the holotype. among killifishes is well-documented (e.g., see RANGE: Known only from the type locality, Parenti, 1981, and references therein). Or- Rio Zapatilla, just south of Lago Grande and estias ascotanensis shares with other species west of Lago Peque-no, in the Titicaca Basin ofthe agassii complex a sexual dichromatism (figs. 22 and 23). described for 0. agassii, above; that is, sub- ETYMOLOGY: The trivial name ctenolepis, adult females and juveniles have a relatively a noun in apposition, from the Latinized mottled pattern, whereas males (and often Greek root ctenos, meaning comb, and lepis, larger females) have a more uniform light to meaning scale, in reference to the presence dark brown pigmentation pattern. In males of ctenoid scales over most of the body of and females, the ventrum is pale yellow to adult males. cream, and otherwise unpigmented. Adults to over 60 mm SL. Orestias ascotanensis also shares with 0. Orestias ascotanensis, new species parinacotensis Arratia and 0. laucaensis Ar- Figure 58 ratia sexual dimorphism with regard to de- DIAGNOSIS: A robust species of the agassii gree of squamation. Females of all three complex, closely related to parinacotensis and species tend to be more fully scaled than 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 197

FIG. 59. Orestias richersoni, new species, Holotype, AMNH 52181, male, 85 mm SL. males; although, the extent of the variation sis to denote the occurrence of this species in among large samples of all three species has Lago Ascot'an, Chile. not been documented. Ventrum and basal portion ofthe pectoral, Orestias richersoni, new species dorsal, and anal fins unscaled. Caudal fin Figure 59 truncate. DIAGNOSIS: A species of the agassii com- Lower jaw upturned slightly more than in plex distinguished by a unique pigmentation other species of the agassii complex, except pattern characterized by a faint to jet black for jussiei and albus, the species with which spot at the dorsal margin of the operculum, specimens now referred to ascotanensis have a black margin of the lower jaw, and a faint been confused (Pellegrin, 1906; Tchernavin, to dark band at the base of the pectoral fin 1944a). rays that extends dorsally onto the lateral Outer jaw teeth unicuspid, tips unpig- shield of scales. Body a deep olive green to mented and protruding beyond epithelium dark brown dorsally, with an iridescent back- covering upper and lower jaws. ground and a faint grayish yellow ventrum. MATERIAL: Holotype (USNM 236790, ex. DESCRIPTION AND REMARKS: Meristic and USNM 167744), an adult female, 54.0 mm morphometric data for this species are sum- SL, collected March, 1919 by W. R. Allen marized in tables 3 through 12. A species of from Lago Ascot'an, Chile. Paratypes (USNM the agassii complex most closely related to 167744, ex. IU 16060), 15 juvenile to adult tschudii, and like it with a somewhat laterally males and females, 36.5 to 53.0 mm, taken compressed body, especially posterior to the along with the holotype; (CAS 46160, ex. IU origin of the dorsal fin. Differing from tschu- 16060), 48 juvenile to adult males and fe- dii meristically by a lower number of verte- males, 22.5 to 62.0 mm, two cleared and brae (32), the higher number (mode of 34, counterstained for bone and cartilage, taken range 33 to 35) autapomorphic for tschudii along with the holotype. (table 3). ADDITIONAL MATERIAL (not part ofthe type Distinguished by its color pattern, the series): Lago Ascotan: FMNH 77384 (160 ju- unique component of which is a faint to jet venile to adult males and females); MNHN black spot at the dorsal margin of the oper- 05.186-188 (three subadults to adults, 33.4- culum. The black margin of the lower jaw 49.3 mm); MCZ 58680 (eight juveniles to and faint to dark band at the base of the adults). pectoral fin rays are present in other species RANGE: Known only from Lago Ascotan of Orestias; however, in no other species (in- (Salar de Ascotan), Chile, a salt basin in the cluding recently collected material) is the pig- northwestern section of Chile (fig. 25). mentation such a deep black, being nearly jet ETYMOLOGY: The trivial name ascotanen- black in most specimens. 198 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 60. Orestias multiporis, new species, Holotype, CAS 46166, female, 85 mm SL.

Adults to 85 mm SL. Males with ctenii on a dorsal profile that slopes gently dorsally from scales posterior to the large and smooth scales the tip of the snout to the anterior base of of the lateral shield. Ventral surface of body the dorsal fin. Distinguished from all other from a vertical through the posterior extent Orestias species by having the median dorsal of pectoral fin to the anal fin bare to fully ridge and lateral series scales, without gran- scaled in some larger adults. Males with ulations, and with multiple neuromasts, oc- breeding tubercles on anal, dorsal and pec- curring in groups of two or more. toral fin rays. Basal portion of pectoral, anal DESCRIPTION: Distinguished from all other and dorsal fin unscaled. Caudal fin truncate. Orestias species by a unique neuromast pat- Outer jaw teeth unicuspid, in two regular tern, characterized by a series oftwo or more or irregular rows, tips pointed and unpig- neuromasts grouped together on each of the mented, projecting beyond epithelium cov- scales of the lateral series and the median ering upper and lower jaws. dorsal ridge. A member of the agassii com- MATERIAL: Holotype (AMNH 52181), an plex, indistinguishable from most other adult male, 85.0 mm SL, collected on Sep- species in the complex in any morphometric tember 4, 1979 by Tom Coon and Pedro or meristic characters (data for this species Quispe, from the Rio Coata, at a bridge just are summarized in tables 3 through 12). upstream from Coata, Lago Titicaca drain- Adults to 85.0 mm SL. All pigmentation age, Peru. Paratypes (AMNH 52182), four patterns characteristic ofPellegrin's varieties subadults to adults, 60.0 to 81.0 mm, taken of 0. agassii present in the type series: ju- along with the holotype. venile males and females and subadult fe- RANGE: Known only from the type locality, males with mottled pattern, subadult and the Rio Coata near Coata, draining into Lago adult males and adult females dark overall, Titicaca, just south of the Capachica Penin- except for ventral surface which is light cream sula (figs. 22 and 23). to yellow in all individuals. Ventrum and ETYMOLOGY: The trivial name, richersoni, basal portion of pectoral and anal fins un- in honor of Peter J. Richerson who with co- scaled, basal portion ofdorsal fin nearly fully workers is carrying out a limnological study scaled. Fully scaled on either side of median of Lago Titicaca; to commemorate my ap- dorsal ridge. All scales smooth, never any preciation for his encouraging a multidisci- granulations, concentric striae present just to plinary and international study of Lago Ti- a limited degree. Males often with breeding ticaca. tubercles on anal and dorsal fins; all fins clear to slightly mottled. Caudal fin truncate. Anal Orestias multiporis, new species pouch small. Figure 60 Outer jaw teeth unicuspid, with pointed cusps, slightly pigmented, arranged in one DIAGNOSIS: A robust species of the agassii scattered row, embedded in epithelium cov- complex, with relatively thick, fleshy lips, or- ering upper and lower jaw margins, tips not bits near the dorsal surface of the head, and visible in most specimens. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 199

FIG. 61. Orestias mundus, new species, Holotype, FMNH 41135, female, 45 mm SL.

REMARKS: In the hypothesis of relation- the complex by being uniformly colored ships within the agassii complex (fig. 30), throughout life, body with a pale yellow ven- multiporis is considered to be in an unre- tral surface becoming light brown dorsally, solved trichotomy with two larger groups of all fins clear or dorsal fin faintly dusky, and agassii complex species; multiporis lacks the a dark thin, midlateral line. defining characters of both groups. Nonethe- DESCRIPTION AND REMARKS: Meristic and less, it is possible that some populations now morphometric data for this species, sum- referred to agassii will be found to be more marized in tables 3 through 12, fail to dis- closely related to multiporis than to any other tinguish Orestias mundus from other species Orestias species. This presumption is based ofthe agassii complex. Nonetheless, it is one on the fact that many collections of agassii of the most distinct and easily identifiable are represented only by juveniles which have species in the complex, differing from the the generally juvenile characteristic of an in- other species by its relatively plain color pat- complete dorsal squamation, but about which tern exhibited with little variation in all adult squamation characters are unknown. known life stages of both males and females. MATERIAL: Holotype (CAS 46166), an adult The absence of marked ontogenetic change female, 85 mm SL, collected November 1918 in pigmentation pattern is characteristic of by W. R. Allen at Maravillas, then a station species of the gilsoni, mulleri, and cuvieri on the southern railway, near Rio de Lampa complexes; however, mundus possesses the (figs. 22 and 23), Peru. Paratypes (CAS defining characters of the agassii complex. 51311), 37 juvenile to adult males and fe- Therefore, this pigmentation pattern is most males, 43-80 mm, taken along with the ho- parsimoniously considered to be autapo- lotype. morphic for mundus. RANGE: Known only from the type locality Adults to 45.0 mm SL. Ventral surface, (fig. 23), on the Rio de Lampa, northwestern basal portions ofpectoral, dorsal and anal fins arm of the Rio Coata (fig. 22), a tributary of unscaled, scales present or absent from either Lago Titicaca, Titicaca Basin. side of median dorsal ridge. Breeding tuber- ETYMOLOGY: The trivial name multiporis, cles present on anal fin of males. Caudal fin a noun in apposition, from the Latin prefix truncate. multi-, meaning many or much, and poris, Outer jaw teeth unicuspid, with pointed plural of the noun porus, meaning a pore or slightly pigmented tips protruding just be- passage, in reference to the multiple series of yond thick epithelium covering upper and neuromasts along the median dorsal ridge and lower jaws. the lateral series. MATERIAL: Holotype (FMNH 41135, ex. FMNH 41130-41140), an adult female, 45.0 Orestias mundus, new species mm SL, collected December 9 to 12, 1941, Figure 61 by C. C. Sanborn, from Cuzco, Peru. Para- DIAGNOSIS: A member of the agassii com- types (FMNH 41130-41134,41136-41140), plex distinguished from all other species in 110 juvenile to adult males and females, 10.0 200 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 62. Orestias ututo, new species, Holotype, AMNH 52183, male, 48 mm SL. to 41.0 mm, one adult cleared and counter- plex, above; that is, a rather mottled pattern, stained for bone and cartilage, taken along often with a more or less distinct midlateral with the holotype. line ofblotches. Adult females with a distinct RANGE: Known only from the type locality, midlateral line, and darkly pigmented dor- Cuzco, Peru in the Urubamba Basin (fig. 24). sally. Adult male holotype (fig. 62) is rather ETYMOLOGY: The trivial name mundus, a uniformly pigmented, with a pale yellow Latin adjective meaning neat or trim, in ref- background and scattered minute dark me- erence to the unadorned, nearly uniform col- lanophores. The color pattern of this adult or pattern from juvenile through adult males male is most similar to the unique pattern and females. diagnostic of Orestias mundus, but differs by being darker overall. The holotype of ututo Orestias ututo, new species is the only large male of the species known; Figure 62 a larger sample size is therefore necessary to determine if there is more variation within DIAGNOSIS: A species of the agassii com- adult pigmentation. plex most closely related to hardini, sharing Ventrum and basal portion of pectoral, with it a low number of dorsal fin rays, the dorsal, and anal fin unscaled; scales on either modal number reduced to 11 in hardini and side of the median dorsal ridge present or 10 in ututo, with an observed range in both absent. Caudal fin truncate. species of from 10 to 12 (table 4). Distin- Outer jaw teeth unicuspid, in one or two guished from hardini and all other Orestias irregular rows; tips black, protruding through by a decrease in the modal number of anal epithelium on upper and lower jaws. fin rays to 11, with an observed range offrom MATERIAL: Holotype (AMNH 52183, ex. 10 to 12 (table 5). AMNH 38416), an adult male, 48.0 mm SL, DESCRIPTION AND REMARKS: Morphomet- collected May 26, 1979 by Tim Hardin, from ric and meristic data for ututo are summa- Lago Ututo, Peru (approx. 9°50'S, 77°30'W). rized in tables 3 through 12. Orestias ututo Paratypes (AMNH 38416), five juvenile to is distinguished from all other Orestias by its adult males and females, 16.4 to 31.0 mm, low number ofanal fin rays (modally 11; ob- taken along with the holotype. served range 10 to 12). ADDITIONAL MATERIAL (not part of type Other characters in tables 3 through 12 do series): cf. ututo: Huayre, Peru (FMNH not serve to distinguish ututo from the av- 70367), 11 juveniles to adults. erage values for agassii complex species; RANGE: Known from Lago Ututo, Peru (fig. however, they do show that it is a relatively 24), north of both the Urubamba and Titi- robust species, deep bodied (table 10) and caca basins, and possibly Huayre, Peru. with a deep caudal peduncle (table 11). ETYMOLOGY: The trivial name, ututo, to Adults to over 40 mm SL. Juveniles with denote the occurrence of this species in Lago the typical color pattern of the agassii com- Ututo, Peru. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 201

FIG. 63. Orestias gilsoni Tchernavin, Lectotype, BMNH 1944.6.6:524, female, 43 mm SL.

Orestias gilsoni Tchernavin ticaca Expedition, at a depth of from 2.5 to Figure 63 2.9 m, off Tiahuanaco. and Orestias gilsoni. Tchemavin, 1944a, pp. 217-219 RANGE: Known from Isla Taquiri off (original description, Isla Taquiri, Lago Pe- Tiahuanaco, Lago Pequefio, Lago Titicaca quefio, Lago Titicaca). (figs. 22 and 23). DIAGNOSIS: A species of the gilsoni com- Orestias taquiri Tchernavin plex most closely related to taquiri with which Figure 64 it shares an increase in the modal number of pectoral fin rays to 17 or 18, with an observed Orestias taquiri. Tchernavin, 1944a, pp. 221-222 range of 17 to 19 (table 6), and a color pattern (original description, Isla Taquiri, Lago Pe- characterized by a silver background with six queino, Lago Titicaca). to nine irregular bars. Orestias gilsoni is dis- DIAGNOSIS AND REMARKS: A species of the tinguished from other members ofthe gilsoni gilsoni complex most closely related to gil- complex by a head that is relatively wide soni, sharing with it an increase in the modal (width reaching nearly 23 percent ofSL; table number of pectoral fin rays to 17 or 18, with 9) and flattened dorsally; and, by a relatively an observed range of from 16 to 18 (table 6), narrow caudal peduncle, its depth usually and a color pattern characterized by a silvery less than 10 percent of SL (table 11). The background with from six to nine irregular combination of these two characters (a wide bars. Distinguished from gilsoni by a reduc- and flat head and a narrow caudal peduncle) tion in the modal number of dorsal fin rays gives gilsoni a distinctive overall body shape. to 12, with an observed range of from 11 to Additional morphometric and meristic data 14 (table 4) as opposed to a modal number for gilsoni are summarized in tables 3 through of 14, with an observed range of from 12 to 12. 15 in gilsoni; and, a reduction in the modal MATERIAL: Lectotype (BMNH 1944.6.6: number of vertebrae to 30, with an observed 524), a female, 43 mm SL, collected July 30, range of from 30 to 31 (table 3), as opposed 1937 by the Percy Sladen Titicaca Expedition to a modal number of 32 vertebrae, with an at a depth of from 2.7 to 2.8 m from Isla observed range of from 31 to 32 in gilsoni. Taquiri, Lago Pequefno, Lago Titicaca, des- Additional meristic and morphometric data ignated herein. Paralectotypes: (BMNH for taquiri are summarized in tables 3 through 1944.6.6:525), a ?male, 28 mm collected with 12. the lectotype; (BMNH 1944.6.6:526-530), MATERIAL: Holotype (BMNH 1944.6.6: eight juvenile to adult males and females, 18 534), an adult female, 26 mm SL, collected to 34 mm; (USNM 133140, ex. BMNH July 30, 1937 by the Percy Sladen Titicaca 1944.6.6:526-530), one adult female, 33 mm, Expedition, at a depth of from 2.7 to 2.8 m, collected with the lectotype; (BMNH from Isla Taquiri, Lago Pequefio, Lago Ti- 1944.6.6:53 1) one adult female, 25 mm col- ticaca. Paratypes: (BMNH 1944.6.6:535, an lected July 28, 1937 by the Percy Sladen Ti- adult male, 23 mm; (BMNH 1944.6.6:536- 202 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 64. Orestias taquiri Tchemavin, Holotype, BMNH 1944.6.6:534, female, 26 mm SL.

542) nine juvenile to adult males and fe- as opposed to 31 vertebrae in tomcooni. Ad- males, one male and one female cleared and ditional morphometric and meristic data are counterstained for bone and cartilage); summarized in tables 3 through 12. (USNM 133141), an adult male, 26 mm, all Tchernavin (1944a) placed this species in taken with the holotype. Group IV, along with mulleri, incae, and RANGE: Known only from Isla Taquiri, one crawfordi because all four species are fully of the two larger islands in Lago Pequei-no, scaled. However, as discussed in the Phylo- Lago Titicaca (figs. 22 and 23). genetic Analysis, the division of Orestias species into fully scaled and unscaled forms Orestias mooni Tchemavin is artificial, and therefore, not the basis on Figure 65 which species complexes are defined in the present study. Lauzanne (1981) describing Orestias mooni. Tchernavin, 1944a, pp. 228-230 ispi, another fully scaled species, comparing (original description, Bahia de Coata in Bahia it with mooni concluded that the two were de Puno, Lago Titicaca). not closely related, a viewpoint that is upheld DIAGNOSIS AND REMARKS: A fully scaled here. species ofthe gilsoni complex closely related MATERIAL: Holotype (BMNH 1944.6.6: to tomcooni, and like it, with a lowerjaw that 503), an adult female, 35 mm SL, collected is at a nearly 90-degree angle to the body axis June 1, 1937, by the Percy Sladen Titicaca (fig. 65). In addition to being fully scaled with Expedition, from Bahia de Coata in Bahia de irregularly distributed small, thin scales, Puno, Lago Titicaca. Paratypes: (BMNH mooni is distinguished by an increase in the 1944.6.6:504-520), 35 juvenile to adult males modal number ofvertebrae to 32 or 33, with and females, 16 to 32 mm; (USNM 133139), an observed range of from 32 to 34 (table 3), an adult female, 28 mm, all taken with the

FIG. 65. Orestias mooni Tchemavin, Holotype, BMNH 1944.6.6:503, female, 35 mm SL. 1 984 PARENTI: ANDEAN KILLIFISH ORESTIAS 203

FIG. 66. Orestias uruni Tchernavin, Lectotype, BMNH 1944.6.6:173, female, 43 mm SL. holotype; (BMNH 1944.6.6:522), two adults pachica Peninsula, Lago Grande, Lago Titi- eviscerated, 24 to 31 mm, collected by the caca, designated herein. Paralectotypes Percy Sladen Titicaca Expedition from Bahia (BMNH 1944.6.6:174-177), four adults, 40 de Taman, Lago Titicaca. to 50 mm, collected with the lectotype. RANGE: Known from Bahia de Coata and RANGE: Known only from the type locality, Bahia de Taman, Lago Titicaca (figs. 22 and Bahia de Uruni on the north side of the Ca- 23). pachica Peninsula, Lago Grande, Lago Titi- caca (figs. 22 and 23). Orestias uruni Tchernavin Figure 66 Orestias minimus Tchernavin Orestias uruni. Tchemnavin, 1 944a, pp. 213-214 Figure 67 (original description, Bahia de Uruni on the Orestias minimus. Tchernavin, 1944a, pp. 216- north side of the Capachica Peninsula, Lago 217 (original description, Molinopampa, on the Grande, Lago Titicaca). southeastern shore of Lago Grande, Lago Titi- caca). DIAGNOSIS AND REMARKS: One of the relatively robust species of the gilsoni complex, DIAGNOSIS AND REMARKS: One of the di- most closely related to tchernavini, robustus minutive species of the gilsoni complex, and imarpe, sharing with them a marbled known only from two adult females, 23 to 28 color pattern. Distinguished from these mm SL, most closely related to the other di- species by the base of the pectoral fin nearly minutive species of the complex, minutus, meeting the posterior margin of the opercu- but distinguished by being totally unscaled, lum, nearly obscuring the basal portion ofthe and by having 30, rather than 31, vertebrae pectoral fin; and, by a reduced or absent anal (table 3). Additional meristic and morpho- pouch and papilla. The polarity of the last metric data for minimus are summarized in character is ambiguous because the anal pouch tables 3 through 12. See also remarks under and papilla are present to varying degrees minutus, below. among most species of the mulleri, gilsoni, MATERIAL: Holotype (BMNH 1944.6.6: and some species of the agassii complexes. 532), an adult female, 23 mm SL, collected However, the absence of these structures in July 16, 1937, by the Percy Sladen Titicaca uruni serves to distinguish it from closest rel- Expedition, at a depth of from 10 to 11 m, atives, and, therefore, at one level, is consid- from Molinopampa, on the southeastern ered to be autapomorphic. Meristic and mor- shore of Lago Grande, Lago Titicaca. Para- phometric characters for uruni are type (BMNH 1944.6.6:533), an adult female, summarized in tables 3 through 12. 28 mm, collected July 13, 1937, by the Percy MATERIAL: Lectotype (BMNH 1944.6.6: Sladen Titicaca Expedition, at a depth offrom 173), an adult female, 43 mm SL, collected 14 to 16 m, from Sucune, southeast of Bahia August 9, 1937, by the Percy Sladen Titicaca de Moho, Lago Grande, Lago Titicaca. Expedition, at a depth of 0.9 m from the RANGE: Known from just two localities in Bahia de Uruni on the north side of the Ca- Lago Grande, Lago Titicaca: Molinopampa, 204 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 67. Orestias minimus Tchernavin, Paratype, BMNH 1944.6.6:533, female, 28 mm SL. on the southeastern shore (figs. 22 and 23), As noted in the explanation of the synapo- and Sucune, southeast ofBahia de Moho (fig. morphy diagrams, both Orestias minutus and 22). minimus are known only from diminutive adult females. Although there are some un- Orestias minutus Tchernavin determined specimens in the collection ofthe Figure 68 AMNH which could be referred to minutus, both species are known with certainty from Orestias minutus. Tchernavin, 1944a, pp. 215- only two specimens each. These specimens, 216 (original description, Bahia de Uruni on the which constitute the type series of each northeastern side of the Capachica Peninsula, Lago Grande, Lago Titicaca). species, are bleached and have fins broken. Nonetheless, each species is distinct and rec- DIAGNOSIS AND REMARKS: A diminutive ognized in this study. Because of the poor species of the gilsoni complex, known from state of preservation, neither of the two syn- two adult females, 24 to 28 mm SL, most types of minutus is chosen as lectotype. closely related to minimus, the other dimin- MATERIAL: Syntypes (BMNH 1944.6.6: utive species in the complex. Defined by an 159-160), two adult females, 24 to 28 mm increase in the number of anal fin rays to 16 SL, collected July 9, 1937 by the Percy Sladen or 17, as opposed to 14 or 15 for minimus. Titicaca Expedition, at a depth of 1.3 m, from Additional meristic and morphometric data Bahia de Uruni on the northeastern side of for minutus are summarized in tables 3 the Capachica Peninsula, Lago Grande, Lago through 12. Titicaca (figs. 22 and 23).

FIG. 68. Orestias minutus Tchernavin, Syntype, BMNH 1944.6.6:159-160, female, 28 mm SL. 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 205

FIG. 69. Orestias tchernavini Lauzanne, Holotype, MNHN 1981:770, female, 46.5 mm SL.

Orestias tchernavini Lauzanne MATERIAL: Holotype (MNHN 1981:770), Figure 69 an adult female, 46.5 mm SL, collected Oc- Orestias tchernavini. Lauzanne, 1981, pp. 84-89 tober 15, 1979 from Estrecho Tiquina, Lago (original description, Estrecho Tiquina, Lago Peque-no, Lago Titicaca. Paratypes (MNHN Pequeno, Lago Titicaca). 1981:771), 36 adult females, 41 to 54.5 mm, taken along with the holotype. DIAGNOSIS AND REMARKS: A species of the RANGE: Known only from the type locality, gilsoni complex most closely related to im- Estrecho Tiquina, Lago Pequenlo, Lago Ti- arpe, uruni, and robustus which together form ticaca. a group distinguished by their overall marbled pigmentation pattern. Orestias tcher- Orestias imarpe, new species navini is distinguished by a relatively laterally Figure 70 compressed head, the width of which is as low as 13 percent of SL (table 9). Additional DIAGNOSIS: One ofthe more robust species morphometric and meristic data for tcher- of the gilsoni complex, distinguished pri- navini are summarized in tables 3 through marily by thick and fleshy lips; a relatively 12, and characters not compared among all small dorsal fin with a modal number of 11 Orestias as part of this study are discussed in rays (observed range of from 10 to 14, table Lauzanne (1981). 4); and large eyes that project above the pri- Tchernavin (1944a, pp. 230-231) gave mary dorsal profile of the head to a greater characters for, but did not formally name, a extent than in robustus, tchernavini, and uru- species that he referred to as "Orestias sp. no. ni, the other robust species within the com- 1." He considered it to be closely related to plex (fig. 70). species in his Group III, which includes DESCRIPTION: Meristic and morphometric species now considered to be part of the gil- data for this species are summarized in tables soni complex. Lauzanne (1981, pp. 88-89) 3 through 12. stated that his new species, tchernavini, was A robust (most closely-related to tcher- conspecific with the specimen referred to by navini, uruni, and robustus) and one of the Tchernavin as sp. no. 1. The specimen larger, species of the gilsoni complex attain- (BMNH 1944.6.6:161) is poorly preserved ing an SL of 57 mm. Orestias imarpe is readi- and I cannot confirm this identification. ly distinguished from these other robust However, I agree with Lauzanne (1981, p. species by its relatively small dorsal fin (rays 89) that tchernavini is rather closely related modally 11, range 10 to 14), and thick and to minimus, gilsoni, and mooni. fleshy lips. Also, the eyes project above the The holotype and 36 paratypes by which primary dorsal profile ofthe head to a greater tchernavini is known are all females. As stat- extent than in other robust Orestias species; ed in Phylogenetic Analysis, female Orestias however, whereas this character helps to sep- are found in much higher proportions than arate imarpe from the other species, it is not males. Whether or not this has any relation considered to be a uniquely derived character to a particular mode of reproduction is un- of imarpe, since the extent of the projection known. ofthe orbits above the primary dorsal profile 206 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 1 78

FIG. 70. Orestias imarpe, new species, Holotype, AMNH 52184, female, 51 mm SL.

of the head varies within species of the mul- 22, 1979, by Tom Coon and assistants near leri and gilsoni complexes. village of Ojjerani, between Puno and Chu- Body with cream-colored background, cuito, Bahia de Puno, Lago Titicaca, at a depth marbled with medium brown reticulations, of from 1 to 2 m at a distance of from 70 to fins dusky or with discrete blotches. Ventral 100 m from the shore. Paratypes: (AMNH surface light yellow. Lower jaw with brown 52185), five adults, 53-57 mm, (one cleared blotches or incomplete band of pigment. and counterstained for bone and cartilage) Marbled pattern characteristic of the group taken with the holotype. including tchernavini, uruni, and robustus as ADDITIONAL MATERIAL (not part of type well as imarpe, faded extensively in alcohol series): Bahia de Puno: BMNH 1982.12.6: in two paratypes that are preserved with 35-36 (two adults, 53-56.1 mm), AMNH mouths wide open. The fading is most likely 52186 (four adults, 50-54 mm); Lago Pe- due to stress of capture and preservation. quefno, near village of Vilurcuni: AMNH Another effect of stress of capture on pig- 52187 (one adult, 56 mm). mentation pattern that can be seen in many RANGE: Known from the type locality in of the specimens collected in 1979 by Tom Bahia de Puno, and from near the village of Coon that has not been described elsewhere Vilurcuni, Lago Pequenlo (figs. 22 and 23). in this paper is the presence of a dark band ETYMOLOGY: The trivial name imarpe, the circumventing the body at a point equal to abbreviated title used by the Instituto del Mar the highest point ofthe dorsal arch. This can del Peru. be seen in the figures ofluteus (fig. 7). In many cases, the dark dorsal band is simply the re- Orestias tomcooni, new species sult ofthe dorsal scale covering being scraped Figures 4B, 18A, 71 off; however, in a large number of lots, the ventral band appears to represent a change DIAGNOSIS: An elongate species of the gil- in pigmentation. This band is presumably soni complex readily distinguished by its caused by stress of capture in experimental overall body shape characterized by a rela- gill nets used extensively by Coon. tively narrow (table 9) and deep head (table Ventrum, basal portion of pectoral, anal, 10), with an even more abrupt transition be- and dorsal fins unscaled; scales of median tween the anterior part of the body and the dorsal ridge small, ridge incomplete in some caudal peduncle than is diagnostic of a specimens. subgroup of the complex comprising gilsoni, Outer jaw teeth unicuspid, with pointed taquiri, minimus, minutus, tomcooni, and tips just protruding beyond epithelium cov- mooni (fig. 71). Modal number of scales in a ering upper and lower jaw margins. lateral series increased to 37, with an ob- MATERIAL: Holotype (AMNH 52184), an served range of from 36 to 38 (table 7). adult female, 51.0 mm SL, collected August DESCRIPTION AND REMARKS: Morphomet- 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 207

FIG. 71. Orestias tomcooni, new species, Holotype, AMNH 52188, female, 53 mm SL. ric and meristic data for this species are sum- of Vilurcuni, on the western side of Lago Pe- marized in tables 3 through 12. Only one such queino. character (modal number ofscales in a lateral RANGE: Both Peruvian and Bolivian shores series increased to 37) distinguishes it from of Lago Pequefno (fig. 22). other gilsoni complex species; nevertheless, ETYMOLOGY: The trivial name tomcooni, it is readily distinguished from all other Or- in honor of Tom Coon with gratitude and estias by its overall body shape, with an abrupt appreciation for his expert collection of more transition between the anterior part of the than 3000 specimens of Orestias from the body and the caudal peduncle. Adults to over Titicaca Basin in 1979, which formed the im- 50 mm SL. Caudal fin truncate. petus for the present revision. Orestias tomcooni is most closely related the con- to 0. mooni, with which it shares Orestias robustus, new species dition ofhaving the lower jaw at a nearly 90- 72 degree angle to the body axis (figs. 65 and Figure 71). In addition to its diagnostic characters, DIAGNOSIS: As the trivial name suggests, tomcooni may be distinguished from mooni one of the more robust species in the genus by its unscaled (rather than fully scaled) ven- Orestias, the width ofthe head reaching near- tral surface and basal portion of the pectoral, ly 23 percent of SL (table 9), and an overall dorsal and anal fins. Also, mooni tends to be body shape characterized by a gentle rise in uniformly straw-colored overall, whereas the dorsal profile from the tip of the snout to tomcooni has a yellow to cream background the base of the dorsal fin, and a gentle down- with mottled lateral and dorsal surfaces. ward slope in the ventral profile from the tip Outer jaw teeth unicuspid with darkly pig- of the lower jaw to the base of the anal fin. mented tips that protrude just beyond epi- Most closely related to a group of species in thelium covering upper and lowerjaws. Teeth the gilsoni complex (uruni, imarpe, and arranged in one or two irregular rows. tchernavini) with which it shares an overall MATERIAL: Holotype (AMNH 52188), an marbled pigmentation pattern. adult female, 53.0 mm SL, collected October DESCRIPTION AND REMARKS: Additional 2, 1979 by Tom Coon and M. Reed, near the morphometric and meristic data for this village of Huatayata, at a distance of from 0 species are summarized in tables 3 through to 30 m from the shore, at a depth of from 12. 0 to 1 m, on the northern side of Lago Pe- Adults to over 50 mm SL. A robust species queino. Paratypes (AMNH 52189) five sub- of the gilsoni complex, distinguished by one adult to adult males and females, 37.0 to 41.0 morphometric character (a greater width of mm, two cleared and counterstained for bone the head) from other species in the complex and cartilage, taken along with the holotype; with which it shares an overall marbled pig- (AMNH 52190) nine juvenile to adult males mentation pattern. and females, 18.0 to 42.0 mm, collected Au- Eyes project just slightly above primary gust 28, 1979 by Tom Coon, near the village dorsal profile; overall body shape robust with 208 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

FIG. 72. Orestias robustus, new species, Holotype, AMNH 52191, female, 45 mm SL. a convex dorsal and convex ventral profile, RANGE: Known from the type locality near contributing to the species' rounded, robust the village ofOjjerani in Bahia de Puno, Lago appearance. Titicaca (figs. 22 and 23) and from north of Ventral surface, and basal portion of pec- the Capachica Peninsula in Lago Grande (fig. toral, dorsal and anal fins unscaled (or basal 22). portion of fins with several scales). Caudal ETYMOLOGY: The trivial name robustus, a fin truncate. Males with ctenoid scales on head Latin adjective, meaning strong or robust, in as well as posteriorly; breeding tubercles on reference to the overall robust appearance of anal fin. Scales present or absent from either this species. side of median dorsal ridge. Outer jaw teeth unicuspid, in one irregular row; tips barely protrude beyond epithelium '?Orestias cuvieri X Orestias pentlandii covering upper and lower jaws. REMARKS: The two species of Orestias that MATERIAL: Holotype (AMNH 52191), an attain the greatest standard length, 0. cuvieri adult female, 45.0 mm SL, collected August (maximum of 220 mm SL recorded) and 0. 24, 1979, by Tom Coon and assistants, at a pentlandii (maximum of 200 mm SL record- depth of from 1 to 2 m, at a distance of from ed) are known from collections throughout 70 to 100 m from shore near the village of Bahia de Puno, as well as other localities in Ojjerani, between Puno and Chucuito, Bahia Lago Titicaca. They were considered by de Puno, Lago Titicaca. Paratypes (AMNH Tchernavin (1944a) to be sister species. The 52192), three adults, 47.0 to 48.0 mm, taken two species are readily distinguished by the with the holotype; (AMNH 52193) eight presence oflarge, recurved outer jaw teeth in specimens, 32 to 48 mm, one cleared and cuvieri, as opposed to few or no teeth in the counterstained for bone and cartilage, col- outer jaws of pentlandii. In addition, cuvieri lected August 21, 1979; (AMNH 52194) two has a larger head, fewer and larger scales in adult females, 48.0 to 57.0 mm, collected Au- a lateral series, and fewer vertebrae than pent- gust 23, 1979; (BMNH 1982.12.6:122-123) landii. With the description of0. ispi by Lau- two adult females, 51.0 mm, collected August zanne (1981), which is hypothesized to be 25, 1979, all from the type locality. most closely related to 0. pentlandii, cuvieri, ADDITIONAL MATERIAL (not part of type and pentlandii are no longer considered to be series): From type locality: (AMNH 52195), sister species (see Phylogenetic Analysis, Ex- one female, 46.0 mm; Escallami, Lago Grande planation of Synapomorphy Diagrams, and north of Capachica Peninsula: (AMNH Systematic Accounts for these three species). 52196), four juvenile to adult males and fe- Tchernavin (1 944a) discussed several males, 25.0-37 mm. specimens collected by the Percy Sladen Ti- 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 209 ticaca Expedition that he considered to be Allen concluded the description by stating hybrids between 0. pentlandii and cuvieri. (p. 381): "Having been unable to find the These specimens (BMNH 1944.6.6:19-20), types, the description is incomplete, and I do two mature males, are intermediate between not know where the species belongs." pentlandii and cuvieri in several characters My own examination of former Indiana including squamation (scales are smaller and University material ofCAS, FMNH, USNM, more numerous than in cuvierl) and head and UMMZ failed to locate the types. There length (head shorter than in cuvieri but longer were numerous collections in the area of Rio than in pentlandii). In both specimens, there Langui by Eigenmann and coworkers in De- is a distinct, large outer row ofrecurved, uni- cember 1918. One lot at CAS, IU 16075, cuspid teeth in the jaws, clearly a character collected on December 8, 1918 from Rio of cuvieri. Body length (165 to 170 mm SL) Langui by C. H. Eigenmann and M. Medina, is within the expected range for males ofboth contains five specimens that are distorted be- cuvieri and pentlandii. yond identification and appear to have been Tchernavin (1 944a, p. 16 1) concluded that dried out at one time. Therefore, Orestias the specimens were most likely hybrids, stat- rospigliosii Eigenmann and Allen must be ing: "Both specimens were taken from among considered a nomen nudum. a school ofripe Orestias cuvieri and probably are the result of interbreeding between this Orestias pentlandii var. fuscus Garman species and Orestias pentlandii." Villwock Orestias pentlandii var. fuscus. Garman, 1895, p. (1964) concurred with this conclusion; how- 148 (nomen nudum). ever, he did not rule out the possibility that these specimens represented males of 0. cu- REMARKS: Garman (1895) concluded his vieri. diagnosis of Orestias pentlandii by stating (p. These specimens cannot be assigned with 148): "Specimens from the Cuzco Valley are certainty to either cuvieri or pent/andii. They about two-thirds as large as those described; may represent a third, undescribed species of they are much darker on the back, above the Orestias, however current collections ofthese lateral line, and apparently belong to a dis- large species are not adequate for such a hy- tinct variety, fuscus." pothesis to be evaluated. No designated type material has been located for this variety; although, there are Orestias rospigliosii Eigenmann and Allen specimens identified as 0. pentlandii from the Cuzco Valley in the MCZ collections Orestias rospigliosii. Eigenmann and Allen, 1942, (MCZ 3941). Garman's sentence does not p. 381 (nomen nudum). serve to sufficiently distinguish fuscus from REMARKS: Allen, in Eigenmann and Allen other forms of Orestias. Therefore, Orestias (1942) published the name of a new species, pentlandii var. fuscus Garman is treated as a Orestias rospigliosii in honor of Dr. Carlos nomen nudum. Rospigliosi y Vigil, former director of the Zoological Museum of the University of San SUMMARY Marcos. The authorship of the species was Forty-three species of Orestias (table 2) are attributed to both Eigenmann and Allen. recognized in the present revision. Fourteen There were to be five specimens, the largest are described as new (gracilis, rotundipinnis, (53 mm total length) the holotype, collected farfani, gymnotus, hardini, ctenolepis, asco- by C. H. Eigenmann, December 8, 1918 from tanenis, richersoni, multiporis, mundus, utu- Rio Langui, Peru, above 12,000 feet eleva- to, imarpe, tomcooni, and robustus). tion. A catalogue number of IU 12290 is giv- Of the 25 forms recognized by Tchernavin en for the type series; however, this number (1944a), two (uyunius and langul) are treated pertains to a lot identified as the characiform as synonyms of agassii, one (0. agassii ow- Curimatella alburnus. The morphometric and enii) is treated as a synonym of jussiei, and other descriptive data are vague and do not one (0. agassii pequeni) is treated as a syn- serve to identify a distinct species of Orestias. onym offrontosus, recognized here as a dis- 210 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 tinct species, although treated by Tchernavin (cuvieri, pentlandii, ispi, and forgeti) defined as a synonym of 0. agassii. The remaining as monophyletic primarily by characters of forms (both species and subspecies) ofTcher- the gill arches. The ventral gill arches are nar- navin are all recognized at the rank ofspecies. row and elongate, and the dorsal gill arches The use of the subspecific rank is abandoned are reduced in size relative to the ventral gill here. Tchernavin's subspecies of 0. agassii arches. The fifth ceratobranchials are closely are judged not closely related (i.e., they do apposed, and are often fused along the mid- not form a monophyletic group). line. Inner teeth are simple and numerous. The genus Orestias is a well-defined mono- Diversity among species of the cuvieri phyletic group. Seven synapomorphies serve, complex reflects the diversity found within as a group of characters, to distinguish the the other three complexes and among the ge- genus from other cyprinodontoid fishes; these nus Orestias as a whole. Two of the species are: (1) the pelvic fins and fin girdle are absent (cuvieri and pentlandii) are large, midwater in all ontogenetic stages examined, (2) the forms of Lago Titicaca; cuvieri has large re- vomer is absent, (3) the middle anal and mid- curved outer teeth and is an omnivore. Or- dle dorsal radials are cartilaginous, rather than estias pentlandii, like its sister species ispi, being ossified, (4) the anterior and posterior has few or no outer teeth and is a planktivore. ceratohyal are separated ventrally by a large The gilsoni complex comprises 10 species gap filled with cartilage, as opposed to being (gilsoni, taquiri, mooni, uruni, minimus, mi- nearly in contact, (5) the first postcleithrum nutus, tchernavini, tomcooni, imarpe, and ro- is absent, (6) the anguloarticular lacks a ven- bustus) defined as a monophyletic group by tral extension parallel to the retroarticular, a derived caudal fin structure. They are all and (7) there are unique squamation and head relatively small, inshore forms found in Lago pore patterns, the latter characterized by a Titicaca. A number ofspecies (e.g., minimus, prominent lyre-shaped arrangement of mi- minutus, and tchernavini) are known only nute neuromasts, the former by a prominent from females. The relatively large sample size dorsal ridge of scales running from the top of of tchernavini (it is known from 37 speci- the head to the anterior base ofthe dorsal fin. mens) suggests that a high proportion of or The neuromast pattern is very similar to all females within species of Orestias may that found in the lowland, tropical South indicate the existence of a particular repro- American killifish genus Cynolebias and has ductive pattern rather than sampling error. been suggested as evidence for the common A high proportion of females is not an un- ancestry of that genus and Orestias. How- common phenomenon within killifishes; the ever, it was shown in a revision of cyprino- extent and nature of the phenomenon needs dontiform fishes (Parenti, 1981) that the two to be investigated further. genera are clearly members of different sub- The mulleri complex comprises five species orders of killifishes, and that this similarity (mulleri, gracilis, crawfordi, tutini, and incae) in head pore pattern should therefore be defined as a monophyletic group by the char- treated as a convergence. acter of having the dorsal as well as the anal In that cyprinodontiform revision it was fin base extending beyond the primary dorsal hypothesized that Orestias has no close South and ventral profile. The extension ofthe anal American relatives, but is in a monophyletic fin beyond the primary ventral profile is one group with the Anatolian cyprinodonts character shared by the mulleri and gilsoni (Aphanius and its relatives), and most closely complexes, which are most closely related. related to the genus Kosswigichthys. Evidence The agassii complex comprises 24 species for this hypothesis is reviewed in the present (agassii, empyraeus, frontosus, polonorum, paper, and is upheld. However, further os- elegans, jussiei, puni, parinacotensis, lau- teological study ofAnatolian cyprinodonts is caensis, tschudii, gymnotus, hardini, cteno- needed to hypothesize the polarity ofa num- lepis, ascotanensis, richersoni, multiporis, ber of characters. mundus, ututo, silustani, luteus, rotundipin- Species of the genus Orestias are divided nis, farfani, albus, and olivaceus). The last into four monophyletic groups, referred to as five comprise what is referred to as the luteus species complexes. group. The cuvieri complex comprises four species All sDecies of the azassii comDlex have a 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 211 series ofenlarged and thickened lateral scales gasta Province, northern Chile. The greatest known as the lateral shield. In a number of concentration of species is found within the species, these scales, as well as the head scales Titicaca Basin, and specifically Lago Titica- are large and smooth, with few or no dis- ca, the large, high-altitude lake of the Alti- cernible concentric striae. In the luteus group, plano of Peru and Bolivia. the head and lateral scales are large and thick, The diversity of Orestias in Titicaca has and covered with granulations. Species ofthe been used as justification for the application luteus group tend to be extremely wide-bod- ofthe term "species flock" to these killifishes. ied and wide-headed. However, the Orestias of Lago Titicaca do Killifishes, as a group, are highly sexually not form a monophyletic group. Therefore, dimorphic and dichromatic. Among Orestias insofar as the concept of a species flock may species, females are generally much larger than be implicit in its monophyly ofincluded taxa, males. All but one species ofthe agassii com- it is recommended that the term not be ap- plex undergo an ontogenetic change in color plied to all Orestias species in Lago Titicaca. pattern that is characteristic of most other The introductions of rainbow and brown killifish. Juveniles and subadult females ex- trout, as well as silversides, into Lago Titi- hibit a mottled pattern. With age, the pattern caca has threatened the endemic Orestias fau- gets denser and adult males and females are na. Orestias cuvieri, the species with large dark on the dorsal and lateral surface, where- adults believed to have been in direct com- as the ventral surface is light. Species within petition with the introduced trout, is pre- the three other complexes, and mundus of sumed extinct. However, reports that Ores- the agassii complex, do not undergo such an tias as a genus is extinct in Titicaca are wrong. ontogenetic change in color pattern, and there Recent collecting has revealed that many is little or no sexual dichromatism. Both ju- species are abundant, and that new species veniles and adults are nearly uniformly pig- remain to be described. Exploration of the mented. many small, isolated lakes of the high Andes The genus Orestias extends throughout the is expected to reveal additional new species Andean Altiplano and beyond, ranging from of this diverse and species-rich genus of kil- Ancash Province, northern Peru to Antofa- lifishes.

LITERATURE CITED Arratia F., G. 1902. Description ofa new cyprinodontid fish 1982. Peces del Altiplano Chileno. In, La ve- from Eastern Peru. Ann. Mag. Nat. Hist., getacion y los vertebrados ectotermicos ser. 7, vol. 10, pp. 153-154. del transecto Arica-Lago Chungarfa. A. Castelnau, F. de Velosa and E. Bustos (eds.). Volumen 1855. Animaux nouveaux ou rares recueillis de Sintesis Proyecto Mab.-6-UNEP pendant l'expedition dans les parties UNESCO (1105-17-01). [only pre-pub- centrales de l'Amerique du Sud, de Rio lication copy seen] de Janeiro a Lima, et de Lima au Para. Arratia, F., G., A. Chang G., S. Menu-Marque, II. Poissons. Paris, xii + 112 pp., pls. 1- and G. Rojas M. 50. 1978. About Bullockia gen. nov., Trichomyc- Cope, E. D. terus mendozensis n. sp. and revision of 1876. Note on the ichthyology of Lake Titi- the family Trichomycteridae. Stud. caca. Jour. Acad. Nat. Sci., Philadel- Neotrop. Fauna Environ., no. 13 (1978), phia, vol. 8, pp. 185-187. pp. 157-194. 1878. Synopsis ofthe cold blooded vertebrata, Bleeker, P. procured by Prof. James Orton during 1859. Enumeratio specierum piscium hucus- his exploration of Peru in 1876-1877. que in Archipelago indico observatar- Proc. Amer. Phil. Soc., vol. 17, pp. 33- um .... Act. Soc. Sc. Indo-Neerl. VI, 49. xxxvi + 276 pp. Cuvier, G., and A. Valenciennes 1860. Ordo Cyprini, Karpers. Act. Soc. Sc. 1846. Histoire des Poissons. Tome 18, Paris, Indo-Neerl. VII (no. 5, deel II), xii + 505 pp. 492 pp. Dingerkus, G., and L. D. Uhler. Boulenger. G. A. 1977. Enzyme clearing of alcian blue stained 212 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

whole small vertebrates for demonstra- and Microcyprini. Proc. Acad. Nat. Sci. tion of cartilage. Stain Tech., vol. 52, Philadelphia, vol. 68, pp. 415-439. no. 4, pp. 229-232. 1940. Zoological results of the second Boliv- Eigenmann, C. H. ian expedition for the Academy of Nat- 1894a. Catalogue offreshwater fishes ofCentral ural Sciences of Philadelphia, 1936- America and Southern Mexico. Proc. 1937. Part 1, Fishes. Proc. Acad. Nat. U.S. Natl. Mus., vol. 16, no. 925, pp. Sci. Philadelphia, vol. 92, pp. 43-103. 53-60. 1943. Zoological results of the second Boliv- 1894b. Notes on some South American fishes. ian expedition for the Academy ofNat- Ann. N.Y. Acad. Sci., vol. 7, art. 5, pp. ural Sciences of Philadelphia, 1936- 625-637. 1937. Part II, Additional new fishes. 1910. Catalogue of the freshwater fishes of Notulae Naturae (Philadelphia), no. 120, tropical and south temperate America. pp. 1-7. Reports Princeton Univ., Exped. Pata- Franz, R., and W. Villwock gonia, 1896-1899, vol. 3, Zool., part 4, 1972. Beitrag zur Kenntnis der Zahnentwick- pp. 375-511. lung bei oviparen Zahnkarpfen der Tri- 1920. On the genera Orestias and Empetrich- bus Aphaniini (Pisces, Cyprinodonti- thys. Copeia, no. 89, pp. 103-106. dae). Mitt. Hamburg zool. Mus. Inst., 1927. The freshwater fishes of Chile. Mem. vol. 68, pp. 135-176. Natl. Acad. Sci., vol. 22, 2nd memoir, Garman, S. W. 80 pp., 16 pls. 1876. I. Fishes and reptiles. In, A. Agassiz and Eigenmann, C. H., and W. R. Allen S. W. Garman, no. 11, Exploration of 1942. Fishes ofwestern South America. I. The Lake Titicaca. Bull. Mus. Comp. Zool. intercordilleran and Amazonian low- Cambridge, vol. 3, pp. 274-278. lands of Peru. II. The High Pampas of 1895. The cyprinodonts. Mem. Mus. Comp. Peru, Bolivia and Northern Chile with Zool. Cambridge, vol. 19, no. 1, pp. 1- a revision ofthe Peruvian Gymnotidae, 179. and of the genus Orestias. Univ. Ken- Gery, J. tucky, xv + 494 pp. 1969. The freshwater fishes ofSouth America, Eigenmann, C. H., and R. S. Eigenmann pp. 828-848. In, E. J. Fittkau, J. Illies, 1892. A catalogue of the freshwater fishes of H. Klinge, G. H. Schwabe, and H. Sioli South America. Proc. U.S. Natl. Mus., (eds.), Biogeography and ecology in vol. 14, no. 842, pp. 1-81. South America. Dr. W. Junk, The Ermin, R. Hague. 1946. Schuppenreduktion bei Zahnkarpfen. Greenwood, P. H. Rev. fac. sci. Univ. Istanbul, ser. B, 1979. Towards a phyletic classification of the Tome 11, fasc. 4, pp. 217-272. 'genus' Haplochromis and related taxa. Everett, G. V. Part I. Bull. British Mus. Nat. Hist. 1971. The rainbow trout ofLake Titicaca (and (Zool.), vol. 35, pp. 265-322. the fisheries of Lake Titicaca). Report Grimm, H. to the Government of the Republic of 1979. Zur Entstehung der Schuppenvariabi- Peru, 180 pp. litiit anatolischer Zahnkarpfen (Pisces, 1973. The rainbow trout Salmo gairdneri Cyprinodontidae). Verh. Dtsch. Zool. (Rich.) fishery of Lake Titicaca. Jour. Ges. (72nd meeting in Regensburg), p. Fish Biol., vol. 5, pp. 429-440. 264. Evermann, B., and L. Radcliffe Gunther, A. 1909. Notes on a cyprinodont (Orestias agas- 1866. A catalogue of the fishes in the British sizii) from central Peru. Proc. Biol. Soc. Museum, London. vol. 6, 368 pp. Washington, vol. 22, pp. 165-170. Hennig, W. 1917. The fishes of the west coast of Peru and 1966. Phylogenetic systematics. Univ. Illinois the Titicaca Basin. Bull. U.S. Natl. Mus., Press, Urbana, 263 pp. vol. 95, 166 pp. Hubbs, C. L., and C. L. Turner Foster, N. R. 1939. Studies of the fishes of the order Cypri- 1967. Trends in the evolution ofreproductive nodontes XVI. A revision ofthe Goode- behavior in killifishes. Stud. Trop. idae. Misc. Publ. Mus. Zool., Univ. Oceanog., vol. 5, pp. 549-566. Michigan, no. 42, pp. 1-80. Fowler, H. W. Humphries, J. M., F. L. Bookstein, B. Chernoff, 1916. Notes on fishes of the orders Haplomi G. R. Smith, R. L. Elder, and S. G. Poss 1984 PARENTI: ANDEAN KILLIFISH ORESTIAS 213

1981. Multivariate discrimination by shape in Altiplano of Peru and Bolivia. Report relation to size. Syst. Zool., vol. 30, no. no. 3, In, H. C. Gilson (ed.), Reports of 3, pp. 291-308. the Percy Sladen Trust Expedition. James, D. E. Trans. Linn. Soc. London, vol. 1, (Ser. 1971. Plate tectonic model for the evolution 3), pp. 27-43. of the central Andes. Geol. Soc. Amer. Myers, G. S. Bull., vol. 82, pp. 3325-3346. 1931. The primary groups of oviparous cy- 1973. The evolution ofthe Andes. Sci. Amer., prinodont fishes. Univ. Ser. Stanford vol. 229, no. 2, pp. 60-69. Univ. Publ. Biol. Sci., vol. 6, no. 3, pp. Konuralp, N. 1-14. 1955. A study on the gonad morphology of Newell, N. D. Anatolian cyprinodontids. Rev. fac. sci. 1949. Geology of the Lake Titicaca region, Univ. Istanbul, ser. B, Tome 20, pp. Peru and Bolivia. Mem. Geol. Soc. 175-199. Amer., no. 36, pp. 1-111. Kosswig, C., and W. Villwock Parenti, L. R. 1964. Das Problem der intralakustrischen 1981. A phylogenetic and biogeographic anal- Speziation im Titicaca und Lanaosee. ysis ofcyprinodontiform fishes (Teleos- Verh. Dtsch. Zool. Ges. Keil 1964. Zool. tei, Atherinomorpha). Bull. Amer. Mus. Anz. Suppl., vol. 28, pp. 95-102. Nat. Hist., vol. 168, art. 4, pp. 335-557. Laba, R. [in press] Biogeography of the Andean Killifish 1979. Fish, peasants, and state bureaucracies, genus Orestias with comments on the the development of Lake Titicaca. species flock concept. In, A. Echelle and Comp. Pol. Stud., vol. 12, no. 3, pp. I. Kornfield (eds.), Evolution of fish 335-361. species flocks. Univ. Maine at Orono Lauzanne, L. Press. 1981. Description de trois Orestias nouveaux Pellegrin, J. du Lac Titicaca, 0. ispi n.sp., 0. forgeti 1904a. Sur les pharyngiens inferieurs chez les n.sp. et 0. tchernavini n.sp. (Pisces, Cy- poissons du genre Orestias. Comptes prinodontidae). Cybium, ser. 3, vol. 5, Rendus Acad. Sci., Paris, vol. 139, pp. no. 3, pp. 71-91. 682-684. Levin, C. B., and N. R. Foster Cytotaxonomic studies in Cyprinodon- 1904b. Mission G. de Crequi-Montfort et E. S& 1972. nechal de la Grange. Note sur les pois- tinae: multiple sex chromosomes in Garmanella pulchra. Notulae Naturae sons des Lacs Titicaca et Poopo. Bull. Soc. Zool. France, vol. 19, pp. 90-96. (Philadelphia), no. 446, 5 pp. Lueken, W. 1906. Poissons, pp. 112-137, In, M. Neveu- 1962. Chromosomenzahl bei Orestias (Pisces, Lemaire. Les lacs des hauts plateaux de Cyprinodontidae). Mitt. Hamburg zool. l'Amerique du Sud. Paris, vi + 197 pp., Mus. Inst., vol. 60, pp. 195-198. 18 pls. Miller, R. R. Reid, G. McG. 1948. The cyprinodont fishes of the Death 1980. 'Explosive speciation' of carps in Lake Valley system of eastern California and Lanao (Philippines)-Fact or Fancy? Southwestern Nevada. Misc. Publ. Mus. Syst. Zool., vol. 29, no. 3, pp. 314-316. Zool. Univ. Michigan, no. 68, pp. 1- Rendahl, H. 155, xv pls. 1937. Einige Fische aus Ecuador und Bolivia. 1956. A new genus and species of cyprino- Ark. Zool. K. Svenska Vetenskapsak- dontid fish from San Luis Potosi, Mex- ad., vol. 29A, no. 11, pp. 1-11. ico, with remarks on the subfamily Cy- Richerson, P. J., C. Widmer, and T. Kittel prinodontinae. Occas. Papers Mus. 1977. The limnology of Lake Titicaca (Peru- Zool., Univ. Michigan, no. 581, pp. 1- Bolivia). A large, high altitude tropical 17. lake. Institute of Ecology Publ. no. 14, Miller, R. R., and V. Walters Univ. of California, Davis, 78 pp. 1972. A new genus ofcyprinodontid fish from Rosen, D. E. Nuevo Leon, Mexico. Contrib. Sci., Nat. 1964. The relationships and taxonomic posi- Hist. Mus., Los Angeles Co., no. 233, tion of the halfbeaks, killifishes, silver- pp. 1-13. sides, and their relatives. Bull. Amer. Moon, H. P. Mus. Nat. Hist., vol. 127, art. 5, pp. 1939. The geology and physiography of the 217-268. 214 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178

Rosen, D. E., and L. R. Parenti 1972. Second discovery of multiple sex chro- 1981. Relationships of Oryzias, and the groups mosomes among fishes. Experientia, vol. of atherinomorph fishes. Amer. Mus. 28, no. 2, pp. 223-225. Novitates, no. 2719, 25 pp. Valenciennes, A. Rowntree, W. S. 1839. Poissons d'Amerique. Rapport sur 1903. On some points in the visceral anatomy quelques poissons d'Amerique rap- ofthe Characinidae, with an enquiry into portes par M. Pentland du lac Titicaca the relations of the ductus pneumaticus et des autres points eleves des Andes. in the Physostomi generally. Trans. L'Institut, ann. 7, no. 276, p. 118. Linn. Soc. London, ser. 2, vol. 9, pp. 47-8 1, pls. 3, 4. Villwock, W. Starks, E. C. 1962. Die Gattung Orestias (Pisces, Micro- 1906. On a collection of fishes made by P. 0. cyprini) und die Frage der intralakus- Simons in Ecuador and Peru. Proc. U.S. trischen Speziation im Titicaca-Seen- Natl. Mus., vol. 30, pp. 761-800, pls. gebiet. Verh. Dtsch. Zool. Ges. (Wien), 65-66. no. 54, pp. 610-624. 1926. Bones of the ethmoid region of the fish 1964. Vermeintliche Artbastarde in der Gat- skull. Univ. Ser. Stanford Univ. Publ. tung Orestias (Pisces, Cyprinodonti- Biol. Sci., vol. 4, no. 3, pp. 139-338. dae). Mitt. Hamburg. zool. Mus. Inst. Stiassny, M. L. J. (Kosswig-Festschrift), pp. 285-291. 1981. The phyletic status of the family Cich- lidae (Pisces, Perciformes): a compara- 1972. Gefahren fur die endemische Fisch- tive anatomical investigation. Nether- fauna durch Einbiirgerungs versuche und lands Jour. Zool., vol. 31, no. 2, pp. Akklimatisation von Freundfischen am 275-314. Beispiel des Titicaca-Sees (Peru/Boliv- Tchemavin, V. V. ia) und des Lanao-Sees (Mindanao/Phi- 1944a. A revision of the subfamily Orestiinae. lippinen). Verh. Internat. Ver. Limnol., Proc. Zool. Soc. London, vol. 114, no. vol. 18, pp. 1227-1234. 9, pp. 140-233. 1976. A contribution to the understanding of 1944b. A revision of some Trichomycterinae the evolution ofmeristic characters, with based on material preserved in the Brit- special reference to Old World cyprin- ish Museum (Natural History). Proc. odontids (Pisces, Cyprinodontidae). Zool. Soc. London, vol. 114, no. 10, pp. Abh. Verh. naturwiss. Ver. Hamburg. 234-275. NF 18/19 (Suppl.), pp. 11-27. 1946. On the lateral line system of some cyprinodonts (Order Microcyprini). Ann. Wiley, E. 0. Mag. Nat. Hist. ser. 1, vol. 13, pp. 429- 1981. Phylogenetics, the theory and practice 432. ofphylogenetic systematics. New York, Terrazas, W. U. John Wiley and Sons, xv + 439 pp. 1970. Lista de peces bolivianos. Acad. Nac. Zufiiga, E. Cienc. Bol. La Paz, 66 pp. 1941. Regimen alimenticio y longitud del tubo Uyeno, T., and R. R. Miller digestive en los peces del genero Ores- 1971. Multiple sex chromosomes in a Mexi- tias. Bol. Mus. Hist. Nat. "Javier Pra- can cyprinodontid fish. Nature, vol.231, do." Year 5, no. 16, pp. 79-86. no. 5305, pp. 452-453.

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